The ABC of Eucheuma Seaplant Production

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Monograph # 1-0703, SuriaLink, July, 2003
The ABC of Eucheuma Seaplant Production

Agronomy, Biology and Crop-handling of Betaphycus, Eucheuma and Kappaphycus the Gelatinae, Spinosum and Cottonii of Commerce
All rights reserved @ 2003 by Iain C. Neish and SuriaLink.com Photos and diagrams by Iain C. Neish except where noted otherwise
1. Abstract & Index
2.Introduction
2.Biology | 3.Agronomy
3.Populations 4.A.TaxonomicClassification | 5.B.TaxonomicDescriptions |6.C.Characteristics | 7.D.TradeNames | 8.E.Tambalang | 9.F.Sacol | 10.G.Spinosum | 11.H.Gelatinae | 12.I.NaturalDistribution | 13.J.CommercialActivity | 14.K.CommercialDistribution |15.L.CommercialSignificance | 16.M.CommercialTrends | 17.N.Morphology | 18.O.LifeHistoryTetraspores | 19.P.LifeHistoryMale&Female | 20.Q.LifeHistoryCarpospore | 21.R.StrainSelection |22.S.VarietalImprovement | 23.T.CarrageenanSynthesis | 24.U.CarrageenanDifferences 4.Locations 25.A.PlantElements | 26.B.GeneralPysiology | 27.C.Temperature | 28.D.Light | 29.E.WaterMotion | 30.F.Salinity | 31.G.Macronutrients | 32.H.Micronutrients&Metabolites | 33.ISiteCharacteristics | 34.J.SitePlacement | 35.K.SiteSelectionTools 36.A.Overview | 37.B.Background | 38.C.HabitatTypes | 39.D.On-bottom | 40.E.Off-bottom | 41.F.PenCageBagTube | 42.G.FloatingLines | 43.H.Rafts | 44.I.PondsRaceways | 45.J.PositionOrientation | 46.K.Attachment | 47.L.Seasonality | 48.M.Startup | 49.N.CroplogBasics | 50.O.CroplogParameters | 51.P.CroplogActions | 52.Q.Bleaching | 53.R.GrazerDamage | 54.S.Grazers | 55.T.AntiGrazer | 56.U.Diseases | 57.V.Weeds | 58.W.GoodAsGold | 59.X.EnvironmentMonitoring | 60.Y.CropMonitoring 61.A.ManagingPropagules | 62.B.TLC | 63.C.MovingPropagules | 64.D.BadWeather | 65.E.Productivity | 66.F.Treating | 67.G.Drying | 68.H.Packing | 69.I.QualityTesting | 70.J.ControlSystems | 71.K.EnviroImpacts | 72.L.Polyculture 8.Acknowledgements 9.Bibliography Abb-Bod|Bor-Dix|Dot-Gle|Gom-Liu|Lux-Ola|Pad-Sid|Smi-Vre|Wal-Zer
5.Methods
6.Crop 7.Glossary
Abstract: The development of commerce based on eucheuma seaplants is an outstanding example of widespread farming that evolved from simple methods refined mainly by farmers in the field. This phenomenon has led to current production exceeding 150,000 dry tons/yr from at least ten countries. Farm-gate revenues on the order of 80-100 M USD remain in rural coastal areas of tropical regions where seaplants are the main source of income to tens of thousands of people. The information base for eucheuma seaplant is expanding but has not yet achieved useful stability. Much "knowledge" about the biology of eucheuma seaplants is practical conjecture extrapolated from studies of other seaweeds. The present monograph is intended as a step toward moving beyond this state of affairs. Eucheuma seaplant production is discussed in light of the plants' biology and the agronomic techniques employed by farmers. Socio-economic factors discussed but emphasis is laid on the characteristics of cultivar populations, factors involving farm location and practical agronomy.
http://www.fishdept.sabah.gov.my/download/ABC_eucheuma_a.pdf; 17/05/2013
Eucheuma seaplant farm, Bohol, Philippines
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The ABC of Eucheuma Seaplant Production all rights reserved @ Iain C. Neish & SuriaLink.com
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Introduction to the Biology of Eucheuma Seaplants

The development of commerce based on eucheuma seaplants is an outstanding instance of how widespread farming of a group of useful plants has evolved from very simple methods that have been refined mainly by farmers in the field.
Farming of the eucheuma seaplants is a continuous process of screening and selection for fast-growing propagules.
Farmers tend to plant and harvest crops on a short cycle of 4-6 weeks. They replant cuttings from the most vigorously growing The eucheuma seaplants, as discussed in the present monograph, plants so reproductive plants tend to be culled out. In past years are defined as being members of the Tribe Eucheumatoideae within spinosum had generally been regarded as being more difficult to the Phylum Rhodophyta, Class Rhodophyceae, Subclass grow than cottonii so there has been much conjecture as to why the Florideophycidae, Order Gigartinales and Family Areschougiaceae. reverse seems true at some sites today. For example there have Commercially the most significant species are Betaphycus gelatinae been suggestions that the commonly cultivated cottonii cultivars ("gelatinae" of the trade), Eucheuma denticulatum ("spinosum" of have been propagated for so many years (e.g. about 30 for the trade) and several species of the genus Kappaphycus ("cottonii" "tambalang") that they are losing vigour while several new spinosum of the trade). The information base for these plants is expanding but cultivars with various national origins been selected from wild stocks has not yet achieved useful stability. Thus much of what is to be said and have reached commercial scales of production. about the biology of the Eucheuma seaplants is practical conjecture, Eucheuma seaplant farming has involved the widespread distribution often extrapolated from knowledge of other seaweeds. The present of cultivars from their source habitats to regions far away from their monograph is intended as a step in the direction of moving beyond origins. this state of affairs. For example Kappaphycus cultivars derived from a few plants The commercial significance of eucheuma seaplants is largely based originating in the Sulu Sea have been distributed to distant seas on their role as raw material for the production of the marine where they form the basis for a commercially significant industry. biopolymer known as carrageenan. There have been not yet been reports of proliferation of natural populations arising from such stocks but the issue of eucheuma Betaphycus spp., Eucheuma spp. and Kappaphycus spp. produce, respectively, carrageenans known commercially as "beta", "iota" and seaweeds as "exotic" or "alien" species has been raised in some jurisdictions. This issue and concerns about general environmental "kappa". The development of commercial cultivation for impacts from seaplant farming are significant concerns to the Kappaphycus and Eucheuma since the mid-1970s has been the major source of expansion for the carrageenan industry and current industry and are addressed in the present monograph. combined production for these seaplants probably exceeds 150,000 Another issue of industry concern is the question of whether "Genetic dry tons per annum at commercial moisture standards of 30-40%. engineering" approaches to cultivar improvement are appropriate for This translates to about 100 M USD worth of dried seaplants and eucheuma seaplants. over 30,000 tons of carrageenan with a value on the order of 250 M Fortunately there are many natural genomes to select from and USD per annum. there are no industry-critical problems that cry out for genetic engineering solutions.
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Introduction to the Agronomy of Eucheuma Seaweeds

The zone of feasible production options for eucheuma seaplant farming is a small subset of all possible options delimited by "crop factors" and by "socio-economic factors" including business, economic and social considerations. The subset of feasible options can be depicted as the "success cube" shown in the paradigm at right.
Success results for a commercial seaplant project only if the best among a wide spectrum of alternatives is implemented.
Note that "success" is defined as being "socially, economically and environmentally sustainable production". The present monograph discusses the biology and agronomy of eucheuma seaplants in light of the three crop factors listed in the the table below. Socio-economic factors are briefly addressed in the present monograph but these will also be the subject of a future monograph that deals with industry structure and the role of strategic business alliances among seaplant-based industries.
Crop factors (i.e. agronomic procedures) critical to commercial success in production systems include: 1. The population, which is the natural population or the population of cultivated seed stocks that is used as the basis of commercial seaplant production. 2. The method, which includes agronomic and processing protocols followed to achieve commercial production within the economic goals set by a given enterprise. 3. The location which must be chosen carefully because good site selection will make or break any commercial seaweed farming enterprise.
Human socio-economic factors germane to the success of commercial seaplant production include: 1. Corporate factors such as structuring of the businesses involved and relations among them. 2. Economic factors including costs, general economic conditions and market conditions. 3. Social factors such as local norms, folkways and mores impacting farm productivity and trading patterns. Socio-economic factors are specific to particular geographic regions. This must be accounted for in agronomic protocol designs.
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3.A Taxonomic Classification

This monograph deals with commercially useful genera of the Tribe Eucheumatoideae; . These are colloquially known as the "eucheuma seaweeds". Their taxonomic classification is : Phylum Rhodophyta, Class Rhodophyceae, Subclass Florideophycidae, Order Gigartinales, Family Areschougiaceae, Tribe Eucheumatoideae The trade names, "gelatinae" "spinosum" and "cottonii" and , generally refer to Eucheuma gelatinae (Esper) J. Agardh Eucheuma denticulatum (Burmann) Collins & Hervey and Kappaphycus alvarezii Doty, respectively. The trade name "cottonii" was originally applied to the wild crop and to the first farmed K. striatum Schmitz. It originally referred to the elkhorn variety but came to be applied to all kappa-carrageenan producing Eucheumatoideae. There is a species K. cottonii but it is a thick, flattened species and has never been farmed successfully.
The trade name "gelatinae", refers to the scientific species Betaphycus (ex. Eucheuma) gelatinae. It may also be applied to any species of Betaphycus that yields beta carrageenan. The trade name "spinosum" generally refers to Eucheuma denticulatum but may be applied to any species of Eucheuma that yields iota carrageenan during extraction. Although the specific name "spinosum" has been used in a taxonomic sense it is applied in this instance as a descriptor of the spiny protuberances typical of the commercial species. The specific name "alvarezii" as applied to the Kappaphycus alvarezii Doty commemorates the late Vicente (Vic) Alvarez, a pioneer in the methods of cottonii agronomy. A good and true friend to many seaplant people including the author (photo right).
Genus: Betaphycus Genus: Eucheuma
Trade name: gelatinae
Symbol: BE
Authority: Doty ex P.C. Silva Symbol: EU
Type species: Betaphycus philippinensis Doty Authority: J. Agardh
Commercial species: gelatinae (GEL)
Common names: Eucheuma, Gelatinae (also see Eucheuma names) Trade name: spinosum
Type species: Eucheuma denticulatum (N.L. Burman) F.S.Collins & Hervey Commercial species: cartilaginium (CAR), denticulatum (DEN), isiforme (ISI), muricatum (MUR) Common names: Agal agal, Agal agal besar, Agar-agar, Agar agar besar, Agar agar pulau, Agar agar seru laut, Chilin-t' sai, Crude agar, East-Indian Carrageen, Eucheuma, Eucheuman, Java agar, Kirinsai, Makassar weed, Ruwe agar, Ryukyu-tsunomata, Singapore weed, Spinosum, Tosaka nori, Zanzibar weed Genus: Kappaphycus Trade name: cottonii Symbol: KA Authority: Doty Type species: n/a Commercial species: alvarezii (ALV), cottonii (COT). inerme (INM), interme (INR), striatum (STT), procrusteanum (PRO) Common names: Agal agal, Agal agal besar, Agar-agar, Agar agar besar, Agar agar pulau, Agar agar seru laut, Chilin-t' sai, Cottonii, Eucheuma, Eucheuman, Guso, Kirinsai
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3.B Taxonomic Descriptions

Eucheumatoideae species of commercial significance are notoriously variable in form. Many cannot be distinguished on the basis of one specimen or collection without further taxonomic study. Since the commercial value of such species is related to the characteristics of the biopolymers that they synthesise the infrared absorption by their gels has come to be a measure of differences among genera and species (Santos, 1989). The generic names "Kappaphycus" and "Betaphycus" reflect the types of carrageenan generally associated with the genera in question.
Species in section Gelatinae tend to grow in very turbulent, active water and seem to be unusually tough. They are compressed, flattened ultimately with noticeable segmentation on some branches. Eucheuma species are conspicuous for their spines.
They often have long, cylindrical branches (e.g. E. denticulatum) or slightly compressed ones (E. serra) . Opposite branches are common. E. serra usually has bilateral spines rather than whorls. It appears to be a shallow-water form that has developed such characteristics in response to bright light and low water motion. For About eighteen to twenty species may be distributed among the four reason of their location, two species, E. isiforme (C. Agardh) J. sections of the genus Eucheuma on the basis of their phylogenetic Agardh and E. uncinatum Setchell & Gardner, are relatively well characteristics (see table). known through the work of Harvey (1853), Dawson (1961), Setchell & Gardner (1924) and a series of more recent investigators including Members of the section Cottoniformia feel different from the rest of the tribe. When alive they are turgid and when bent they will snap or D. P. Cheney, C. J. Dawes, James N. Norris and P. W. Gabrielson. These species generally have odd life cycles and their spines tend to break like a fresh carrot. When dry they are without spines and be irregularly scattered. There are other less well-known members of somewhat ropey. Very dry material may be covered with salt (KCl) this section that are very different in form. Doty (pers comm.) crystals. When one looks at individual thalli, be it on herbarium suggested that they should be set off in discrete subsections. The paper or alive in the field, the occasional thallus is hard to place. Among Kappaphycus species young primary branches generally turn cystocarps are borne subterminally on special branchlets or determinate spines and consequently these species often bear spines upward and are relatively blunt, especially in young specimens of K. striatum. Secondary branches may start as hemispherical bumps and asymmetrically. eventually become spiny. If branches become spiny they tend to turn Eucheuma section Anaxiferae holds the species Eucheuma arnoldii downward but do not form whorls. Branching is never truly opposite Weber-van Bosse and E. amakusaensis Okamura. and the branches may be in pinnate series in part of a thallus, especially on long branches. New branches often arise from the dimly These are distinguished by a lack of differentiation in the central axial regions of their segments and by indeterminate vegetative lit interior of dense thalli, then grow toward the light, especially growth of the sexually fertile branches. These are initially spine-like among cultivated plants growing on suspended lines. On the other and produce cystocarps subterminally. This indeterminate growth hand, some species may be strongly compressed. Doty (1988) and appears to bring mature cystocarps to the surface of mature Santos (1989) treat this section as the distinct genus Kappaphycus segments. Possibly, as in E. isiforme (Gabrielson, 1983), it is Doty. initiated in subapical papillae.
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3.C Table of Eucheumatoideae Characteristics

Characteristics of the typical species in the major sections of Eucheuma within the Tribe Eucheumatoideae: Cottoniformia and Eucheuma. Eucheuma section Cottoniformia 1- Fronds of many forms but commonly cylindrical; simple blunt or spiny protuberances 2- Protuberances irregularly arranged; in some cases appearing in vertical rows so segments become angular 3- Branching irregular; sometimes irregularly pinnate; may be opposite or falsely dichotomous 4- Hyphal axial core usually present; not rhizoidal; cylindrical. 5- Kappa carrageenan 6- Cystocarps on main axes 7- No cystocarp associated with laterals Eucheuma section Eucheuma 1- Fronds cylindrical; spines simple; basal diameters less than axis thicknesses. 2- Spines in regularly spaced pairs or whorls first, but later others may appear scattered 3- Branches generally from whorls; often opposite; irregular or in pectinate series 4- Axial core rhizoidal and cylindrical 5- Iota carrageenan 6- Cystocarps on lateral axes 7- Generally a terminal spine beyond cystocarp
Characteristics of the typical species in the major sections of Eucheuma within the Tribe Eucheumatoideae: Gelatinformia and Anaxiferae. Eucheuma section Gelatiformia 1- Fronds compressed; spines simple; basal diameters equal axis margin thickness 2- Spines in rows, marginally first and latter dorsally and ventrally on flatter faces or scattered 3- Branches mostly marginal, pinnate, often opposite or irregular but not in pectinate series 4- Axial core tortuous, hyphal and often flattened. 5- Beta or other carrageenans 6- Cystocarps on lateral or pedical axes 7- Often several spines on cystocarp; sometimes none Eucheuma section Anaxiferae 1- Fronds cylindrical or dorsoventral and bearing compound spines 2- Spines often scattered, in whorls or covering the thallus in various arrangements or densities 3- Branching generally from whorls; often opposite; whorled or irregular 4- No hyphal or rhizoidalaxial core in cylindrical axes 5- Iota carrageenan 6- Cystocarps on main axes 7- Ultimately no spine on cystocarp
Note: These descriptions were provided to the author by Maxwell S. Doty during discussions held in Honolulu on several occasions in the early 1990s. The author bears any responsibility for errors or omissions. The author welcomes amendments, additions and updates from those willing to contribute.
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3.D Trade Names and Varieties

Since eucheuma seaplant farming commenced after 1970 there has been wide dissemination of strains around the world and a proliferation of cultivars as well; especially with Kappaphycus Occasionally strains appear that have poor processing characteristics and these are usually eliminated from contention fairly quickly. In other cases there are mixed reactions to strain quality. Since the Philippines has hosted commercial farming for the longest time it is there that the greatest number of Kappaphycus variants seem to have arisen. At least two variants of kappa carrageenan are found among these; the K. alvarezii type with a distinct infrared absorption peak at wave no. 805 and the K. cottonii types that lack this peak (Doty & Santos, 1978). Most cultured strains are of the former type but Aguilan et al (2003) have recently reported that the Sacol strain is of the latter type. The following table describes Kappaphycus strains now in commerce but this list is not comprehensive and the proliferation of strains continues. 1. Tambalang type: long strands; typically fewer branches than flower; small to large diameter branches; generally thriving in deep water in more northern areas of the Philippines but seldom seen in the Southern Philippines as "flower" now predominates there. Also the predominant strain in much of Indonesia, India, Sabah, Malaysia and Tanzania. 2. Flower type: short strands; bundles of multiple branches resembling a "flower". Found in shallow reefs areas of the Philippines. Dominant strain in the Bongao/Sitangkai areas of the Philippines since 2000. Also seems to be appearing in South Sulawesi and Nusa Tenggara Timur in Indonesia. 3. Vanguard type: Shorter than tambalang but bigger than flower. Found in farm areas of southwest Mindanao.
4. Bisaya type: Looks like a cross between tambalang and sacol types. Predominant form in the Bohol region of the Philippines. 5. Sacol type: clumps of short multiple branches, with small diameter stems. Often found over sandy or muddy substrate such as that found near its source area of Sacol Island, Zamboanga, Philippines. Sold as salad vegetable in Cebu markets. This strain is recently being replaced by farmers with the "Bisaya" type. 6. Sumba type: Long, thick strands. Rather like a coarse, robust form of the tambalang type. Originated in Sumba Island, Indonesia but now grown at several sites in Indonesia. Favoured by some farmers in Bali.
Aside from the Sacol type all of these appear to be strains of Kappaphycus alvarezii. As comparisons of DNA characteristics are extended to strains their relationships should become more clear. For example Aguilan et al (2003) used such methods to compare various strains of K. alvarezii, Kappaphycus sp. sacol variety and Eucheuma denticulatum. Different strains of Kappaphycus alvarezii appeared to have similar banding patterns regardless of their differences in morphology and habit but Kappaphycus sp. "sacol" variety from Bohol showed a different banding pattern. The sacol type may have closer affinity to K. cottonii that to K. alvarezii.
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3.E Kappaphycus alvarezii var. tambalang (cottonii of the trade)

Kappaphycus alvarezii var. tambalang plant (lower left) and closer view of branch (upper right) showing a cylindrical axis with branches that are commonly enlarged maximally just beyond the basal curvature toward the light as manifested through the "candelabra effect" (see below). Clean, upswept tips such as these are typical of healthy, rapidly growing plants. This sweep of fronds toward the light results in a candelabralike appearance
M.S. Doty drawings; I.C. Neish photos - Length of bar ca. 10 cm. per image lower left
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3.F Kappaphycus sp. var. sacol (Sacol cottonii - may be new species)
Kappaphycus sp. var. sacol (Sacol Island strain; shown right) is one of several cultivars that appears to have been propagated by vegetative propagation from plants obtained from wild stocks. The morphology of the eucheuma seaplants tends to be quite variable and can result from genetic differences among strains; environmental factors; agronomy methods; and apparently from spontaneous mutations that occur within a strain and lead to sustained characteristics such as colour differences. The light brown plants in the foreground, below, are examples of Eucheuma denticulatum but the darker brown plants (left), the green plants (middle rear) and dark red plants (back, right) are all thought to be Kappaphycus alvarezii var. tambalang variants.
Taxonomic classification of Kappaphycus species based on morphology is notoriously difficult difficult because of the extreme plasticity of this genus. Recently published data suggest that based on molecular analysis using the rbcL the Kappaphycus sp. Sacol variety is most likely a form of K. cottonii (Aquilan et al, 2003).
M.S. Doty drawing - Length of bar ca. 10 cm. relative to image
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3. G Eucheuma denticulatum (spinosum of the trade)

Eucheuma denticulatum is less variable than Kappapycus spp. among sites and types. The plants generally appear as shown here except that the colour may vary from light brown to deep read (almost black); the branches may be more or less spindly; and the density of "spines" may range from sparse to dense. It is thought that several indigenous varieties have been developed from local wild stocks in the Philippines, Indonesia and Tanzania. There has been some dispersal of these stocks; notably with the dispersion of Bali spinosum to the Central Philippines.
Drawing I.C. Neish after M.S. Doty - Length of bar ca. 10 cm. relative to image
Appearance aside, one distinguishing characteristic of E. denticulatum is a distinct "chlorine" (probably bromine) odour that becomes especially noticeable during drying.
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3.H Betaphycus gelatinae (gelatinae of the trade)

Betaphycus gelatinae is cultivated and/or harvested from wild stocks to a much lesser extent that Kappaphycus or Eucheuma species. It is the smallest of the eucheuma seaplants and it tends to inhabit sites with active water motion. Fronds are apically flat, pliable and arising from marginal cylindrical teeth.
M.S. Doty image - Length of bar ca. 10 cm. The carrageenan from Betaphycus is of commercial interest and is rather close to agar and furcellaran in its applications performance. These small plants are awkward to handle using conventional eucheuma seaweed cultivation methods and fast growing varieties do not seem to have been developed so Betaphycus farming has not yet achieved significant commercial proportions. Some has been cultivated in China both on monolines and by attaching propagules to stones or coral using string or elastic bands (photo right).
Gelatinae drying in Hainan, China
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3.I Natural Distribution and Dispersion by Humans

Natural stocks of Eucheuma seaplant species occur naturally throughout the Indo-Pacific region from eastern Africa to Guam. They are most commonly found between about 20 degrees north and south of the equator in the Indo-Pacific and this zone is roughly defined by the winter isoclines of 21 and 24 degrees Celsius (Doty, 1987). The greatest abundance of these species seems to be in the algal reef areas of island archipelagos associated with Southeast Asia. They generally grow interspersed with corals and at first glance can often be mistaken for corals. There are also outlying species of relatively localized distribution. Three northern outliers are Eucheuma uncinatum in the Gulf of California, E. isiforme in the Caribbean and E. amakusaensis in southern Japan. There is also E. deformans from Lord Howe Isl., and E. speciosum (Sonder) J. Agardh in southwestern Australia as well as E. platycladum (Schmitz) and E. odontophorum (Boergesen) in Tanzania and Mauritius, respectively. Doty (pers. comm) suggested the possibility of biogeographic distribution changing in response to crustal changes in the earth. The species in Australia, aside from those at its most northward edge, are mostly unique and seem to have been developed with odd mixtures of genomes from species further north. In this regard one may list Eucheuma deformans and E. speciosum. One would note that there appear to be unlabeled specimens in herbaria that do not fit into the specific concepts ordinarily recognized among commercial species (Doty, 1988). Eucheuma serra is found nested well within the borders of the distribution of E. denticulatum and Kappaphycus alvarezii is within the distributional borders of K. striatum.
Human actions have had a major impact on the distribution and abundance of eucheuma seaplants. For example Kappaphycus alvarezii seems to have been narrowly restricted to the southernmost Sulu Archipelago, the Celebes Sea and Biak na Belau north of the equator until after 1974 it became widely distributed by man. The occurrence of "cottonii" in Ponape before 1971 may be an introduction from further west during Japanese occupation of the area. Kappaphycus striatum has been taken to Japan recently (Mairh et. al., 1986) and K. alvarezii has been taken to India Mairh et al (1995). K. alvarezii and E. denticulatum have both been taken to Hawaii where, although they not abundant, they are classified as "alien and invasive" algae. From Hawaii they have been taken to eastern and western Kiribati, Tonga, Fiji and elsewhere (e.g., to the Society Islands and temperate North America). While Eucheuma isiforme were also brought to Hawaii along with Hypnea musciformis by some businessmen. From the Philippines both K. alvarezii and E. denticulatum strains have been taken to the Lombok Straits area of Indonesia and have since spread throughout Indonesia. E. denticulatum has also spread via Singapore to Djbouti. K. alvarezii of Philippine origin forms much of the basis for cottonii farming in Indonesia but the substantial spinosum production of Indonesia is based primarily on strains originating from Bali. Balinese plants have since spread to Sabah and the Philippines and seem to be widely cultivated in those countries now. Although The Philippine K. alvarezii has been the basis of most cottonii farming throughout the world. there are also local strains which have been commercially grown in Indonesia, Tanzania and Malaysia.
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3.J Commercial Activity

Where eucheuma seaplants are transferred among sites the live material has generally been shipped in plastic bags or jute sacks wet enough to prevent desiccation. Sometimes transport has taken several days and a high proportion of the transported material has died. The author is aware of instances (e.g. in Tanzania and Bali) where surviving material amounting to only tens of grams has led to commercial farming. Today propagules are routinely shipped in multi-ton quantities from nursery areas to farm areas. This is usually done by loosely packing live plants in woven cloth that are kept shaded and are frequently wetted.
Commercial production of "cottonii of the trade" is currently on the order of 80-120,000 dry tons per year at a commercial standard of 38% moisture-content. The available figures indicate that about 99% of cottonii is commercially cultivated in four countries. Relative production volumes are approximately as shown in the Table 3.K with the Philippines at 70%, Indonesia at 24%, Malaysia (Sabah) at 4%, Tanzania at 1% and others (e.g. Fiji, Kiribati) at a total of 1%. Experimental farming or intermittent commercial activity has occurred in several countries including China, Japan, Ponape, the Solomon Islands, the USA (Hawaii), Belize, Maldives, Cuba, Venezuela Vietnam and India. In all of these countries cultivation has utilised Kappaphycus cultivars of Philippine origin. In some cases (e.g. the "Sumba strain" in Indonesia) local cultivars have also given rise to commercially useful cultivars but it appears that a substantial percentage of the world crop is still descended from material that initially developed to commercial scale in the Philippines. The demand for "spinosum" is less than 1/4 the demand of "cottonii" so the world production of Eucheuma spp. seems to be on the order of 20-25,000 tons. During the early years of cultivation development Eucheuma was more difficult to grow than Kappaphycus and it used to command a higher price. In recent years farmers have found Eucheuma to be the more easily grown and the market has had a persistent oversupply. Betaphycus is a slow growing genus that has yet to make a serious market impact, although there have been serious efforts to find and propagate fast growing varieties (e.g. by the author).
Several countries now produce significant amounts of commercially dry eucheuma seaplants (moisture content generally 30-45% and averaging about 38%). Accurate statistics are unavailable since secrecy and obfuscation are commercial devices still prevalent in the trade. However estimates can be made based on the apparent volumes of the commercial trade. Estimates presented here are drawn mainly from the SuriaLink's trade contacts.
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3.K Commercial Distribution
Distribution of commercial and development activity for eucheuma seaplants. Production in dry tons per annum (% in brackets). ( * Production figures from SuriaLink.com; ** Legend: +++ = large; ++ = medium; +=small; - = none ? = in doubt) '000 km. Kappaphycus (world %) Year Year Production of coast Introduced commercial circa 2002* 15.3 (1.81) 1985 2000 800 (0.7) 7.0 (0.83) 1989 2002 200 (0.2) 54.7 (6.48) 1975 1986 48,000 (42.0) 4.8 (0.57) 1998 1999 300 (0.3) 4.7 (0.55) 1977 1989 4,000 (3.5) 36.3 (4.30) 1971 1973 60,000 (52.5) 1.4 (0.17) 1989 1991 1,000 (0.9) 114,300 (100)
ca. 1,400 386 7.5 (0.89) 0.4 (<0.1) 3.8 (0.44) 0.3 (<0.1) 1.1 (0.13) 0.1 (<0.1) 2.5 (0.29) 0.1 (<0.1) 0.8 (<0.1) 29.8 (29.75) 0.5 (<0.1) 1.1 (0.13) 0.6 (<0.1) 9,330 6.1 (0.72) 2,470 1.9 (0.22) 0.4 (<0.1) 5.5 (0.65) 0.4 (<0.1) 0.1 (<0.1) 0.2 (<0.1) 2.8 (0.33) 3.4 (0.40) ca.1980s ca.1980s 1995 2000 1991 1973 ca.1978 1978 ca.1985 ca.1985 1978 1983 1996 1977 1986 ca. 1980s ca.1985 ca. 1990s 2000 ca.1978 1987 1983 1977 1971 1996 1993 not known not known pre-comm. pre-comm. pre-comm. ceased pre-comm. ceased ceased ceased ceased ceased pre-comm. pre-comm. pre-comm. pre-comm. pre-comm. pre-comm. pre-comm. ceased pre-comm. pre-comm. pre-comm. ceased pre-comm. pre-comm. nil trace small/variable small/variable small/variable nil small/variable nil nil nil nil nil small/variable small/variable small/variable nil small/variable trace small/variable ceased small/variable small/variable small/variable nil small/variable small/variable
Country China India Indonesia Madagascar Malaysia Philippines Tanzania Totals

Antigua, Barbados, Jamaica, St. Lucia Belize Brazil Cambodia Cuba Djibouti Fiji French Antilles French Polynesia Guam Honduras Japan Kenya Kiribati Maldives Mexico Micronesia (Ponape) Mozambique Myanmar Samoa Solomon Islands Tonga Tuvalu USA (Hawaii) Venezuela Vietnam
Eucheuma Production circa 2002* nil nil 8,000 (35.7) 400 (1.8) trace 10,000 (44.6) 4,000 (17.9) 22,400 (100)
trace trace nil nil nil nil nil nil nil nil nil nil trace nil nil trace nil trace nil nil nil nil nil nil nil nil
Develop Status **Expand 2003 potential Sources expand + Wu et. al. (1988) expand ++ Mairh et. al. (1995), Neish (obs.) expand +++ Adnan & Porse (1987), Neish (obs.) expand ++ Ask & Corrales (2002) expand ++ Neish (obs.), Doty (1980) contract ++ Doty & Alvarez (1973), Parker (1974) static + Lirasan & Twide (1993)
static ceased static expand static ceased static ceased ceased ceased ceased ceased static static static unknown static unknown expand ceased expand static static ceased static expand + + ? + ? ? + + + + + + + + + + ? + + + ? + de Boer, pers.comm.; Zemke-White, in press de Boer, pers.comm. de Paula et al. (1998, 1999) Pers. Obs. of author Serpa-Madrigal et al. (1997) Braud & Perez (1978) Prakash (1990), Luxton et al. (1987) Barbaroux et.al. (1984) Doty (1985a) Doty (1985a) Neish (obs.) Mairh et al. (1986) M. Fazal, (pers. Comm.) Luxton & Luxton (1999) de Reviers (1989) Robledo, 1998 Doty (1985a) Fazal, pers.comm. Pers. Obs. of author Doty (1978a) Tanaka (1990), Smith (1990) Tanaka (1990), Fa'anunu (1990) Gentle (1990) Doty (1985a,b) Rincones & Rubio (1999) Ohno et al. (1995, 1996)
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3.L Commercial Significance
Figure 3.M. Graphic History of Carrageenan Seaweed Production; 1961-2002 A. Estimated production of the commercially cultivated warm water seaweeds Kappaphycus spp. In Malaysia (MY), Indonesia (RI) and the Philippines (RP) 1975-2002. (from SuriaLink.com & Seaweed Industry Association of the Philippines (SIAP)). B. Estimated production of commercially harvested wild cold water red seaweeds (mostly Chondrus crispus; some Furcellaria fastigiata) from France (FRA), Canada (CAN) and the U.S.A 1961-2001. (from SuriaLink.com after FAO and Prince Edward Island Fisheries Dept. statistics)
This figure gives an overview of general trends that have taken place with respect to the commercial production of some major carrageenan seaweeds (carrageenophytes) during the past four decades. The producing countries taken into account are those that produce most of the carrageenophytes in their respective segments.
Note: Landings of wild carrageenan bearing seaweeds in Chile fluctuated between about 7,000 13,000 tons/annum and averaged about 10,000 tons/annum from 1991 1999. There was no indication of declining harvest. Genera harvested were Mastocarpus, Sarcothalia, Gigartina and Chondracanthus. (Avila & Pavez, 2000)
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3.M Commercial Production Trends 1961-2002

The graph for tropical production that is presented in section 3.L is based on production of Kappaphycus spp. ("cottonii"). The production of Eucheuma spp. ("spinosum") occurred more or less in parallel and production has generally run about 20% of Kappaphycus production; about 2/3 from the Kappaphycus producing companies noted on Figure 3.L and 1/3 from Tanzania (see photo below).
Comprehensive statistics are available for all years and all locations but the numbers below appear to reflect the following industry trends: 1. Since reaching peak levels on the order of 40-60,000 tons per annum in the decade from 1967-1977 The Chondrus harvest has shown a steady decline to the point where it has leveled off at about 6-10,000 tons per annum. 2. Although this decline may be partly attributable to declining effort industry sources suggest that many populations of Chondrus have diminished in size and/or that the distribution of harvestable beds has become reduced. 3. In areas such as Prince Edward Island in Canada there has been a tendency for Chondrus and Furcellaria to grow in mixed populations. This reduces crop value as plants must be separated to facilitate extraction. 4. During the period 1977 - 1991 average Chondrus and Furcellaria production held steady at about 25,000 tons per annum before declining to present levels around 1992.
Cold water seaplants such as Chondrus (right) can be cultivated but production costs are generally too high for economic production of carrageenan raw materials.
5. Commercial cultivation of cold water carrageenophytes has been technically successful but is economically attractive only for highvalue crops such as specialty foods. 6. By 1977 commercial cultivation of eucheuma seaplants became firmly established and growth of the carrageenan industry was driven by the availability of cultivated tropical carrageenophytes. 7. One major development precipitated by the availability of cultivated Eucheuma seaweeds was the development of "Processed Eucheuma Seaweeds" (PES - E407a) which was cost effective and opened new carrageenan markets. 8. The development of cultivation in tropical Asia is causing a trend for carrageenan process capacity to migrate from Europe and North America to Asia.
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3.N Morphology
The kappacarrageenan-bearing Kappaphycus species referred to as "cottonii" exhibit a highly variable gross morphology. It is difficult, if not impossible, to distinguish species among herbarium materials. (Kappaphycus examples shown right)
Rhizoids in E. denticulatum dominate the central axial region. Small cells may be conspicuously mixed with large cells, depending on the species. Thylles arise by budding yeast-like from large medullary or inner cortical cells and persist as small, somewhat elongated cells among large ones, especially in the central axial region. Kappaphycus displays no abundance of such small cells in its mature axial tissues (Weber-van Bosse, 1913). Among iota-carrageenan producing species those in the section Anaxiferae similarly have a paucity of small cells among the large ones seen in an axis.
The internal vegetative morphology of the eucheuma seaplants appears to be based on variations in the the patterns of cells arising from longitudinal axial filaments (Doty 1985; Doty & Norris, 1985). In sections Cottoniformia and Anaxiferae the longitudinal axial filaments are soon transformed into somewhat elongated large round cells. In sections Eucheuma and Gelatiformia the fate of the long rhizoids occupying the axial cores must be different but just how is not yet clear.
Generally K. cottonii and K. procrusteanum are compressed and clearly distinct due to the dorsoventral flattening and generally prostrate habit of the former and the generally erect, flat, orbicularbladed nature of the latter.
The walls of eucheuma seaplant cells notoriously thicken with age and the relationships between them become hard to trace, though The tissues of Eucheuma comprise an outer cortex of radial filaments, there are "pits" between them. a subcortical/medullary area and a central core. Methods have been developed for precisely identifying where The outer cortex may contain as few as four cells in tetrasporic thalli different carrageenans are located in the cell walls of eucheuma seaplants and these may lead to further elucidation of the biological (Doty, 1988). On the other hand there may be very many cells as in vegetative portions of Eucheuma speciosum (Harvey 1853). Generally role of carrageenan and the structure of cell matrices (Vreeland et al. 1987; Zablackis et al., 1988). By such methods, florescin has been the subcortical/medullary area is formed of large, thick-walled cells. conjugated (Vreeland et al., 1987) to carrageenan and the molecules Below the apex in true K. cottonii the axial region is made up of are labeled when potassium is adequately present. relatively large cells that are not readily distinguishable from peripheral medullary cells. In most other species a core can be distinguished. It is made apparent by the presence of small cells. In kappa-carrageenan producing species such as K. alvarezii these cells may be the thylles of Weber-van Bosse (1926).
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3.O Life History - Overview & Tetrasporangia

The life cycle of eucheuma seaplants is not well known. Perhaps the most significant agronomic breakthrough during the development of eucheuma seaplant cultivation was in the realization that one need not go through spore production in order to propagate crops. Vegetative clones proved to grow indefinitely and some varieties have now been maintained in this manner for more than thirty years. Eucheuma and Kappaphycus are considered to have a triphasic life cycle with gametophyte (N), tetrasporophyte (2N) and carposporpophyte (2N) phases. Evidence seen in some species and the sparse knowledge of male sexual thalli suggests the possibility of other life histories Since E. uncinatum is a seasonal species and the reproduction method is unknown for several other species of seasonal and/or restricted distribution it seems likely there are deviants from the ideal triphasic life cycle (Norris, 1985; Dawson, 1961; Cheney, 1975; Kraft, 1969; Azanza-Corrales, 1990). The reproductive contents of Eucheuma tetrasporangia always become seriately arranged. Mature tetraspores seen in situ can be separated from each other and appear to be without walls. Yokochi (1983) studied the formation of tetrasporangia in B. gelatinae and found that liberation of their spores occurs during the warm season. There is a tendency for terminal spores to remain less than cylindrical and before discharge these are not broadly rounded even when nearly mature (e.g., Kraft, 1969, 1972; Gabrielson, 1983; Doty, 1988; Azanza-Corrales, 1990). Eucheuma uncinatum appears to have terminal spores so reduced in size that they are presumed to be sterile. The ontogeny of tetrasporangia and tetraspores is harder to follow than one might suppose despite their large size and abundance. The most mature sporangia appear to be subterminally pitted. Their cell walls are hard to see by the usual staining techniques and seem to disappear though hydration into gels that lead to spore discharge.
Living tetrasporophytes readily discharge spores which, when caught on slides in the laboratory, are individually spherical and separate from each other. Those of Kappaphycus alvarezii readily adhere and grow on a variety of substrata. However their development has been seen in few species since laboratory facilities are scarce where Eucheuma is common. During his detailed description of B. gelatinae tetraspore discharge and subsequent development Yokochi (1983) observed that spore discharge peaked in November. The tetraspores were reported to be only 15-25 m in diameter upon discharge. The smaller erect germination stages are shown as bearing hairs. The largest were smooth and hairless.
M.S. Doty image
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3.P Life History - Male and Female

Male structures are seldom seen among eucheuma seaplants except as reported in Eucheuma isiforme. According to Cheney (1975) and Dawes et al. (1974: 244f), they appear in superficial sori apparently on thallus segments. AzanzaCorrales (1990) has also observed this in Kappaphycus alvarezii. This is typical for the family. The latter authors apparently found male structures in the proportions to be anticipated by simple genetic sex segregation. During earlier studies Kraft (1969) illustrated that these common but infrequently reported cortical structures were articulated hairs. These structures are not yet generally accepted as male structures by phycologists. The female reproductive morphology of eucheuma seaplants is also little known. Kraft (1969) made a significant advance in describing the carpospores as solitary. In K. procrusteanum Kraft he distinguished sterile suspending filaments between the fertile gonimoblast filaments. Cheney (1975) illustrates the carpogonial branch of E. isiforme. The reproductive morphology of that species was later treated in detail by Gabrielson (1983). Further details concerning eucheuma seaplant reproductive structures were reported for E. serra, a close relative of the commercial "spinosum" by Gabrielson & Kraft (1984). Furthermore the work of Norris (1985), Doty (1988) and (Azanza-Corrales, 1990) add various details about K. alvarezii reproductive structures. From such work it is apparent that the female structures of eucheuma seaplants are typically in spherical structures called cystocarps. These are readily visible as hemispherical to spherical bodies arising from the thallus surface. The distinctive morphogenesis of apical tip growth of E. isiforme has been described by Gabrielson (1983) and this morphogenesis, along with that to be inferred from Gabrielson & Cheney (1987), appears to set Eucheuma apart from closely associated genera of the Solieriaceae.
In the Eucheumatoideae fertile spine development is important systematically (Doty, 1988). In Eucheuma section Anaxiferae spines bear cystocarps subapically. These grow into branches of vegetative form. In other species they are not known to do so. Fertile spines in some species of section Eucheuma are morphologically distinct. They elongate and bear several cystocarps whilst remaining very slender. They appear to be sessile on the main branch axes as do those of Eucheuma section Anaxiferae but the derivation is different. In Eucheuma speciosum tetrasporangia are borne in what appear to be deciduous special branchlets that may be rejuvenated (Doty, 1988). On the assumption that these propagate and the compound spiny nature of the cystocarps of E. speciosum is advanced Doty considered these two species to be among either the most highly- or the least-derived in the genus.
M.S. Doty image
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3.Q Life History - Carpospores

There is little published on the shedding of live carpospores and their development into fertile thalli (Azanza and Aliaza, 1999).
The placenta includes filaments that radiate between the vegetative outer lining of the cystocarp lumen and the fusion cell (e.g., Okamura Carposporangium walls appear to dissipate around the forming spore. 1906,1936, Plate 276; Weber-van Bosse 1928; Kraft 1969). This gelation of diploid gonimoblast cell walls is not understood but From a detailed study of Eucheuma isiforme, Gabrielson (1983) may be the source of the force separating and pushing the mature suggested that these filaments arise from the gonimoblasts and grow spores from the cystocarp. When they are shed carpospores will to the pericarp. However in some species they seem to pass through attach to glass and can be reared in dishes. The most complete the concentric gel-filled space independent of the gonimoblast description is for the infrequently seen species Eucheuma filaments and suspend the spherical fusion cell, abetting its centric amakusaensis (Shinmura, 1975). Shinmura records the discharge of position and radial development. In Kappaphycus alvarezii these carpospores and illustrates their development to the point where they suspending filaments (Doty, 1988) appear to be present before, and form an elongated cylinder about 0.4 mm tall. In K. alvarezii each independent of, the branched gonimoblast filaments and may have a diploid embryo of this type becomes a multicellular ball with rhizoids. different origin. Later the top protrudes so that juvenile thalli become pear-shaped and appear to be covered with a thick layer of gel through which long unstructured hairs are seen to radiate and protrude (Doty 1987). Laboratory conditions for K. alvarezii carpospore shedding have recently been studied by Azanza & Aliaza (1999) who showed the potential of using spores for farming of this species. Within cystocarps the diploid spores, called carpospores, arise from gonimoblast filaments. These in turn radiate from a central fusion cell into a gel layer bounded outwardly by vegetative haploid cells and thence to the vegetative pericarp wall of the cystocarp. This feature found, for example, among farmed representatives of Kappaphycus alvarezii (Azanza-Corrales, 1990). Doty (1988) observed only one cystocarpic specimen of E. gelatinae but on this specimen in some cases a cystocarp was borne on a morphologically distinctive re-branched spine. If one follows the terminology of Harvey (1853) the cavity of the cystocarp may be spoken of as a placenta.
M.S. Doty image
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3.R Selection of Cultivar Strains

It seems that most, if not all commercially cultured strains of eucheuma seaplants have been obtained by the continuous selection that occurs as farmers plant and harvest their crops more or less on a daily basis.
Generally speaking farmers select plants on the basis of their growth rate; indeed the process of long-term culling of vegetative plants leads inevitably toward such selection. In some cases, however, aesthetic attributes (e.g. colour preference by farmers) may have played a part in strain selection.
One of the first and most dramatic instances of field selection was when screening in the Sulu Sea led to discovery of the species that came to be recognised as K. alvarezii var. tambalang (Doty, 1985). This strain has become dispersed to several parts of the world and is thought to be the predominant commercial strain. Since 1971 several genetically stable variants of K. alvarezii have come to be farmed commercially. The most common are known as the green, olive-green, red and brown types. Each strain has found favour in several farming areas and all four are roughly equivalent in the quantity and quality of the carrageenan that they yield.
Above: Colour variants in the harvest at Gunung Payong, Bali, Indonesia Left: Tending the crop in Kudat, Sabah, Malaysia.
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3.S Varietal Improvement

To date genetic work by classic hybridisation techniques has not led to varieties of eucheuma seaplants superior in growth, farmability or carrageenan characteristics. The lack of widely recognisable male individuals and the large size of the thalli may prevent use of the conventional Mendelian genetic breeding methods used successfully by Van der Meer (e.g. 1979) in Gracilaria and by Fang et al. (1963) and Fang (1983) in Laminaria. Cheney (1986) and Polne-Fuller & Gibor (1987) have developed methodologies of protoplast fusion. Cheney (pers. comm.) also reports on a technique called "cell-cell" fusion which has been developed to genetically modify strains of commercially valuable red seaweeds including Eucheuma. Their efforts have been toward removing the cell walls from around protoplasts without damaging the cytoplast. Techniques to establish culture procedures that will result in a large number of cultivars and inexpensive propagation of Kappaphycus have been presented. For example Dawes and Koch (Azanza, pers. comm.) developed procedures for micropropagule and tissue culture to develop propagules successfully introduced to farms in the Philippines.
Recently Reddy et al (2003) in India have utilised in vitro somatic embryogenesis and the regeneration of somatic embryos to yield whole plants. The generation of propagules was successfully demonstrated when pigmented uniseriate filamentous calli of Kappaphycus alvarezii (Doty) Doty were utilised as the basis for axenic cultures. More than 80% of the explants that they cultured using solidified Provasoli enriched seawater (PES) medium showed callus development. Excised calli grew well in subcultures and maintained prolonged growth if transferred to fresh medium in regular intervals. Propagule production was further improved through somatic embryogenesis by culturing thin slices of pigmented callus using appropriate media. Transfer of embryogenic calli along with somatic embryos to agitated liquid media facilitated rapid growth and yielded propagules that grew into whole plants during subsequent cultivation in the sea. The daily growth rate of one tissue cultured plant was monitored for seven generations in field and found to be as high as 1.51.8 times the rate of farmed plants propagated through vegetative means alone. The suggest their techniques as a tool for rapid and mass clonal production of seed stock of Kappaphycus for commercial farming.
There is a school of thought that advocates "Genetic engineering" approaches to cultivar improvement among for use with eucheuma seaplants but this is a controversial issue. Detractors of this view point out that market resistance can be expected as a result of an expanding global movement, against genetically modified organisms (GMO) and many people value seaplants and their products largely because of their "naturalness". Some major users of carrageenan stipulate that GMO seaplants must not be used as raw material for manufacture of their ingredients. Fortunately there are still many natural gene pools to select from and there are no major problems with eucheuma seaplants that cry out for genetic engineering solutions. There is still plenty of scope for programs of screening for new vigorous strains derived from natural populations of Kappaphycus and retention of "seed banks" for strains of interest.
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3.T Carrageenan Synthesis

Eucheuma seaplants are eaten as sea vegetables but their main commercial value is due to the carrageenan that is a structural component in their cell walls.
Carrageenan is synthesised in the Golgi apparatus. Enzymatic Carrageenans are complex polysaccharides considered to be processes defined by the plants' genetic codes effect its structure (e.g. pattern of sulphation). Synthesised carrageenan is exported via molecular hybrids of several basic units (G4S-DA, G0S-DA, G4Scisternae to the cell wall and excreted to the matrix. DA2S, etc.). Genes determine the presence of the biosynthetic enzymes involved in carrageenan synthesis but the actual process is not a template mechanism that makes perfect copies as with proteins under DNA or RNA control. Kappaphycus will always produce kappa carrageenan but environmental parameters have effects on molecular structure. The plants have a natural growth, maturation and senility cycle that influences carrageenan composition. Azanza-Corrales & Saa (1990) have shown there is seasonal variation in the gel quality in K. alvarezii. The gel from K. alvarezii improves in strength with the diameter of the thalli (i.e. with age) but it appears that both haploid and diploid plants have similar carrageenans. This is in contrast to Chondrus and various other Gigartinales. There is still ongoing experimentation to find ways to get more and better gels from K. alvarezii and E. denticulatum in the face of opposing interests of farmers and extractors. Farmers seek methods that involve less work or provide more weight (be it water, sand or seaweed) for they sells by weight, not by yield or quality. On the other hand extractors wants fewer tons of lessexpensive seaweed with higher yields and more valuable gel per ton. This can lead to some "to-and-fro" in agronomy practices. For example during early farm development pruning of growing tips was practiced but this led to a preponderance of low gel strength young material being marketed and such methods are now discouraged. Kappaphycus kappa carrageenan comprises alternating units of 3-linked D galactosyl (G unit) and 4-linked D galactosyl (D unit). The G unit is mostly sulphated at carbon 4 (G4S). The D units are often sulfated at carbon 6 (D6S).
Seaweed buyers generally encourage harvesting methods that yield predominantly older plants in order to obtain higher gel strength or higher viscosity gels. Optimum harvest time is 6-8 weeks after planting (Barraca, 1989).
The D units may also be converted to 3,6 anhydro galactosyl sugar (DA). Variations to these basic units include a G unit that is unsulphated (G) and a D unit sulphated at carbon 2 (DA2S).The D6S sugar is considered as the biological precursor of the DA sugar. This conversion can also be achieved by chemical methods (e.g. alkaline modification). Carrageenan molecules are flexible and provide fibre components and/or a matrix in which skeletal fibres are embedded. In function it may be analogous to more rigid glucans such as cellulose, mannans and xylans that occur in other seaplants and land plants.
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3.U Carrageenan Differences Among Species

Carrageenans are characteristically sulphated with the exception of "beta" which lacks sulphation and therefore strains the definition of "carrageenan" (Santos, 1989). Greer and Yaphe (1984) considered beta-carrageenan to be a precursor of gamma-, mu- and kappa-carrageenan but it appears in high concentration with some kappa-carrageenan in B. gelatinae and B. speciosum. Kappa carrageenan from K. procrusteanum and K. cottonii lack infrared absorption at Wave Number 805 (Doty & Santos, 1978) and this differentiates their carrageenan from the kappa-carrageenan of K. alvarezii (Santos, 1989). Aguilan et al (2003) have recently shown that such is also the case with the "Sacol" strain of Kappaphycus and this, along with DNA evidence, may lead to re-classification of the Sacol variety. The lack of a peak at the 805 position is interpreted as being due to a complete lack of sulfate in the 2-positions of the galactose units. Today farmed material is thought to be predominantly K. alvarezii so when one says "cottonii" the reference is usually to varieties of that species. Strictly speaking, though, the commercial term "cottonii" may include any or all species of the section Cottoniformia. Carrageenan from the more slowly growing K. striatum and E. inerma Schmitz do show infrared absorption at Wave Number 805, which means that at least some of the galactose moieties are sulfated at their 2-positions. Some species such as E. odontophorum and E. platycladum are reported to contain both iota and kappa carrageenans. (Mshigeni & Semsi, 1977; Santos, 1989). Thus far such species have not attracted significant commercial interest.
Iota-carrageenan is produced by all species in Eucheuma section Eucheuma which contains E. denticulatum and in section Anaxiferae which contains E. arnoldii and E. amakusaensis. The iota-carrageenans from the species E. uncinatum and E. isiforme are deviant types and less commercially useful than E. denticulatum (Lawson et al., 1973; Penman & Rees, 1973; Dawes, 1979). Another factor bearing on commercial value is that E. isiforme is not known to be present in quantities sufficient to supply commercial needs. Laboratory gel-yields are often double those obtained in commercial extractions. Most extractors must be content with yields of as low as 25 percent. In some cases laboratory yields from cottonii have run over 75 percent. Yield may be lower with age in cottonii thalli but the strength of the gels made using carrageenan extracted from older Kappaphycus thalli appears to be greater. Since cottonii is mostly prized for gel strength farming so that the average age of the harvest is older is advised. The generally accepted practise in most commercial farming areas is to harvest after the crop about 6-8 weeks after planting. Gel strength may even be high in branches displaying the symptom of malaise called ice-ice so in harvesting or buying such bleached segments of thalli should not be discarded if the interest is gel strength. However, it was shown that low quality crops characterized by low carrageenan yields and gel strength are due to the state of crops during harvest (Trono and Lluisma 1992). In slow growing and "sickly" crops (with ice-ice) the amount of pure carrageenan yield is 25 to 40% less than those from healthy crops (Trono 1993).
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4.A
Basic Elements of Seaplant Growth
Agronomy methods must be developed with a clear understanding of what seaplants need in order to grow Note that plants differ from animals in aquaculture in these important ways: 1. They get all their nutrients from the seawater that moves past them 2. They must absorb photons in order to photosynthesize and grow 3. They utilize carbon dioxide and generate oxygen; the opposite of animals 4 Nitrogen compounds of the type excreted by animals are crucial nutrients to seaweeds 5. Unlike mobile animals such as fish, seaplants have no means of suspending themselves in the water column These factors have important implications for the types of culture systems which must be used for raising seaweeds; including: 1. Enclosures or suspension systems must expose the plants to both light and water flow so largevolume enclosures such as salmon cages cannot be used 2. Plants must be separated and suspended by the use of physical structures (e.g. cages or ties on ropes) and/or applied force (e.g. air and paddlewheel agitation). 3. Plants and animals can be combined in properly designed polyculture systems to their mutual benefit The best fertiliser is a farmers shadow on the field This is a time-tested principle of terrestrial agronomy that also applies to seaplant farming.
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4.B
General Physiology
There is little physiological work published on eucheuma seaplants. Since eucheuma seaplants are of commercial interest some studies have been made to elucidate relationships between the environment and seaweed production but much basic biology remains to be done. Consequently many of the principals of eucheuma seaweed physiology and biology must be extrapolated from results obtained during studies of other seaplants and land plants. General trends expected on that basis are as follows: Anabolic processes including the rate of photosynthesis and growth rate generally follow a pattern such as the one illustrated at left. These processes respond to environmental variables such as temperature, light intensity and the availability of essential nutrients. By this model anabolic processes increase as the environmental variable increases, reach a "Peak" or a "plateau" where conditions are best and diminish once the variable exceeds the optimum range. At some point death occurs.
Catabolic processes such as respiration generally respond to environmental variables such as temperature according to a pattern such as the one shown here. By this model catabolic processes increase as the environmental variable increases, reach a "plateau" around the level at which conditions are best for plant growth and increase once the variable exceeds the optimum range. At that point catabolic losses increasingly offset anabolic gains. At some point tissue death occurs.
It is known that some carrageenan bearing seaplants such as Chondrus crispus tend so store essential nutrients such as nitrogen once nutrient levels exceed the quantity needed for growth. If nutrient levels drop below the level necessary for growth the plants will maintain growth and other anabolic processes by utilising this stored nitrogen. within ranges typically found in seawater levels of essential nutrients such as nitrogen probably do not reach levels that harm the plants.
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4.C
Temperature Responses
Temperature affects can directly effect the plants' physiological processes or they may indirectly effect plants by temperature impacts on the surrounding environment. For example temperature effects water motion, hence farm productivity, by generating wind, waves and currents.
Species Kappaphycus alvarezii Eucheuma amakusaensis Eucheuma denticulatum Eucheuma isiforme Kappaphycus striatum Eucheuma uncinatum Max/range in deg. C 22.8 - 29.2 18 - 25 Max. 34 Max.18-25 20 to 30; 22.8-29.2 Max. 24 Author or source Ohno & Orosco (1987). Ohmi & Shinmura (1976). Dawes (1979). Mathieson & Dawes (1974) Mairh et al. (1986); Ohmi & Orosco (1987). Dawes (1979).
In Southeast Asian farm areas sea temperatures do not often reach critical levels but optimum ranges have been observed. Although farmers report rapid growth and high biomass production by Kappaphycus during months characterised by water temperatures ranging between 25C and 30C (Barraca, pers. comm.). Njoman et al. (1987) found temperatures of 27 to 32C where they were rearing K. alvarezii on the south-western shore of Sumatra. Ohno & Orosco (1987).
The photosynthesis and respiration rates of eucheuma seaplants seem to be significantly influenced by temperature. Glenn & Doty (1981; see table 4.C above) found physiological maxima for photosynthesis at 25C for three species and they also observed that respiration increased 50-60% from 15C to 20C. Auto-oxidation increased sharply from 25C to 40C. Dawes (1979) found E. isiforme to have a respirometric maximum at about 3040C. E. uncinatum and E. denticulatum had maxima near 30C. For E. isiforme photosynthesis reached a maxima of 30-40C depending on temperature (Mathieson and Dawes, 1974). Dawes (1979) reported that when thalli were returned to a standard 25C after maxima were tolerated the values previously obtained at 25C were not repeated consistently. He interpreted this as evidence of tissue damage at high temperature. Field studies indicated that E. isiforme, began to grow seasonally in March and continued until after August when there was a general decline in correlation with temperature and other seasonal factors.
Above: Perennial Philippine Eucheuma farms seen from ca. 100 m.
In higher latitudes eucheuma seaplants may be sensitive to seasonal cold and may even disappear during winter. For example the strong seasonality of E. uncinatum is well known. This species is confined to the Gulf of California. It appears in spring and dies off in the summer in response to temperatures that exhibit wide seasonal variation. In culture Zertuche et al. (1988) observed negative weight changes in winter when temperatures were at their lowest. Another "northward outlier", E. amakusaensis, grows with the spring increase in temperature until early summer (Ohmi & Shinmura, 1976) and by late summer it disappears (Shinmura, 1975). Ohno & Orosco (1987) and Mairh et al. (1986) reported on Kappaphycus striatum transferred in 1983 to Tosa Bay, Japan from the central Philippines. It was studied in field culture but in November, when temperatures fell below 20C, they found that the plants died.
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4.D Light Responses

Exposure to optimal amounts and wavelengths of photosynthetically active radiation (PAR) is undoubtedly as essential for eucheuma seaplants as it is for other plants. Mairh (1986) found that with K. striatum maximum growth rates were obtained with a 12:12 L:D cycle at 6,000 lx. but that growth rates started to drop above about 10,000 lx. Dawes (1979) reported that growth rates increased to about 18,000 mW cm-2 of white light for E. denticulatum, E. isiforme and E. uncinatum. Eucheuma appears to have a daily photosynthetic rhythm both for photosynthesis and respiration, (Glenn & Doty 1981). It appears that Eucheuma is opportunistic with respect to light insofar as photosynthesis is concerned. Results with Kappaphycus were similar. Photosynthesis in Kappaphycus shows a diurnal pattern with a peak in late morning. Glenn & Doty (1981) observed that storage conditions could effect such periodicity. During storage trials there was strong periodicity from the thalli stored overnight outdoors and strong suppression of periodicity in thalli stored indoors. It appears that excessive light can have deleterious effects on the growth of eucheuma seaplants. This has been attributed to excessive light in respect to other elements in the environment. In non-tidal situations such as on floating rafts or in shallow ponds; or in sites with high reflectance from white sandy bottoms sudden crop death has been noted and this seems be light-related. In early farming days Doty noted (pers. comm.) that by June 22 secondary branchlets of K. striatum turned away from the light and by late May or June many thalli died on the shallow reef flats then in use. These could be offset by moving thalli to bottoms of greater water depth. Zertuche et al. (1988) obtained best growth by shading culture tanks to achieve light intensities measured where the thalli were collected in the wild.
Commercial eucheuma seaplants are generally grown in suspended culture within one meter of the sea surface or on structures attached to the sea floor where water depth is about one meter at low tide. On some farms the adverse effects of more intensive sunlight near the surface of the water column can be offset by intensive planting such that distances between lines and cuttings are close enough to permit shading (e.g. Trono (1994). Another method that works is to move plantings to a greater depth. When a farmer has succeeded in placing plants close to optimal conditions eucheuma seaplants tend to exhibit a growth habit sometimes referred to as a "candelabra effect". In such cases plants are very clean, have even pigmentation and have profuse branches distinctly growing toward the light. There may be differences among eucheuma seaplant strains with respect to photosynthetic responses. Dawes (1992) found that different colour variants of K. alvarezii and E. denticulatum exhibited different responses. Such responses may also be effected by the abundance of nitrogen storage compounds in plants since many such storage products are present in the form of pigments. As a general rule it appears that the darker the plant, the more stored nitrogen it contains. A single plant may exhibit a range of colour spanning from very pale yellow to very dark brown or green over the course of several weeks and this can be expected to have an impact on photosynthetic responses.
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4.E Water Motion

It has generally been noted that eucheuma seaplants grow best in moving water. Water motion helps to clean plants, bring fresh nutrients, remove metabolites and apply hydraulic forces that stimulate plant growth.
Within the physical breakage limits of the seedlings and equipment one can approximate that faster water flow equals faster growth. The author has observed instances (e.g. in San Bernardino Strait in the Philippines) where rapidly flowing water produced Kappaphycus plants as much as two meters long and with major branches more than two cm across (photo opposite).
One important factor concerning water motion effects is the unmixed boundary layer. The thickness of this layer is inversely proportional to water movement and turbulent flow disrupts the boundary layer. Still water can expose plants to low nutrient availability and high waste levels.
The unmixed boundary layer is caused by friction between the thallus and surrounding waters. Eucheuma seaplants generally grow on algal reef flats and tidal movements are important in structuring their habitats. As the tides periodically subside desiccation occurs and the sun's radiation results in a decapitation of biological reef growths. This leaves level reef flats near the extreme low water mark. This combination of causes results in the flat non-consolidated, relatively level, and uniform environments where both cottonii and spinosum tend to grow in the wild.
The unmixed boundary layer appears to be one factor that causes water motion to effect eucheuma seaplant growth but perhaps physical stresses also stimulate growth. Effects of water motion are confounded with those of temperature, light and nutrients. Effects of the sun and the moon influence the water motion so important to Kappaphycus growth. In the field both the moon-driven tides and currents generated as a result of solar heating provide water motion. The sun's heat causes both wind (hence waves) and oceanic currents. When the water is deepest (e.g., at high tide) flow tends to be laminar. At the lowest low tides water motion tends to be turbulent. The erect Kappaphycus species stand rigidly in place whether growing from immobile rock or tied on a farm with water moving turbulently through them and enhancing inward and outward materials diffusion.
Azanza-Corrales et al. (1995) showed the importance of water motion in the "seeding" or natural sporulation experiments done in Eucheuma and Kappaphycus farms in the Philippines.
Water motion is an important factor to take into account during the selection of farm sites and during crop logging. See section 5 of the present monograph for procedures.
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4.F Salinity and Water Quality

The effects of nutrition and salinity on Kappaphycus are not well known although it can be assumed that they are of critical importance to plant growth. Kappaphycus seems to grow best in "full salinity" seawater. At most successful farm sites salinity seems to be on the order of 30-35 ppt. Dawes (1979) found E. isiforme to have a broad respirometric maximum at about 30-40 ppt. Both E. uncinatum and E. denticulatum had maxima near 30.ppt. Salinity and terrestrial influences are significant for seaplants in both negative and positive ways.
Table 4.F shows nutrient analyses of Hawaiian waters with ocean surface values for comparison (g-atm l). Codes: "D" (dipped), "P" (pumped from bay or shore), "W" (pumped from wells) or, in reference to Keahole Point water provided by the National Energy Laboratory Hawaii, "S" (shallow, warm) or "C" (deep, cold). (Doty et al. 1986). Water source Salinity ppt. 24 15 34 1 5 32 34 PO4 NO2+NO3 NH4 Si
1- Anuenue Fish. Res. Ctr. "W" 1.1 15.9 36.0 193.7 2- Aurea Marine Inc., Kahuku "W" 1.7 7.8 64.6 178.7 3- MRTC pond Kaneohe Bay "P" 3.2 23.4 4.2 45.1 Seawater strongly influenced by land is significantly different in 4- Lilipuna Pier, Kaneohe Bay "D" 0.1 0.1 2.1 28.7 salinity and micronutrient content from pristine open-ocean water 5- Waiahole stream water "D" 0.1 0 0 27.8 such as that which usually bathes wild eucheuma seaplant 6- Waiahole + Wahiawa soil 0.3 1.3 0.1 23.5 populations (see table). 7- Coconut Is., Kaneohe Bay "P" 18.5 0.7 0.9 6.0 Water from wells and near-shore areas have a high silica content but 8- North reef of same Island "D" 0.1 0.2 1.4 5.6 it may be 0.1 g-atm l or less in ocean water. 9- Keahole warm NELH 20 m "S" 0.2 0.2 0.4 3.0 Note the wide variation in both fixed nitrogen and phosphate values 10- Natatorium reef Waikiki"D" 0.1 0.1 0.1 1.9 and the ratios between them (Table 4.F opposite). Though it has not 11- Keahole cold NELH 610m "C" 3.0 39.0 0.2 0.2 been possible to date to obtain analyses of the water surrounding 12- Ocean surface <2.9 1-50 <5-50 <0.7-1.8 wild eucheuma seaplant populations they probably receive water near the lowest silica values such as the oceanic surface water shown (Lines 11 and 12). In water favorable to other seaweeds some Eucheuma species such Glenn and Doty's (1990) 55-week study in Hawaii utilized Coconut as Eucheuma denticulatum may die. Island North Reef water (Line 8) in the field. In Anuenue water (Line 1) E. denticulatum always died quickly and In laboratory work they usually used either Natatorium Reef (Line disappeared, as did E. isiforme when out-planted in water from 10) or Anuenue Fisheries Research Center water (Line 1). The best Kaneohe Bay (Line 8). In terrestrially contaminated well waters K. growth of Eucheuma spores was obtained in such water or in water striatum lingered on and K. alvarezii grew at about half its usual from Kaneohe Bay (Lines 4 & 8). The shallow ponds mentioned rate. above under "Light" had Coconut Island water flowing into them (Line 7).
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4.G Macronutrients
Among macronutrients it seems generally accepted that nitrogen is crucial for productive farming and that this nutrient can be a limiting factor in sea cultures. In Hawaii macronutrient conditions were measured upstream and downstream of an unusually dense planting of eucheuma seaplants (Glenn & Doty, 1990 & Table 4.G below). The results indicated utilization of phosphates, nitrate and nitrite and production of ammonium nitrogen as water flowed through the plantings. The net production of ammonia was attributed to resident grazer populations. Sample site Upstream Downstream Water motion NH3-N NO3+NO2-N PO4-P Oxygen 45 38 1.98 2.01 1.44 1.10 0.66 0.62 7.22 8.25
A bleaching phenomenon similar to the "Neish Effect" is observed among eucheuma seaplants (see photo below) Thus far, however, adequate coupled laboratory activity is not known. Zertuche-Gonzales et al. (1993) showed that higher carrageenan yields were obtained from cultures with fertilized E. uncinatum under high light conditions.
Pending further work with eucheuma seaplants perhaps one may be forgiven for postulating that have nitrogen metabolism fundamentally similar to that of Chondrus. The work of Neish and Shacklock (1971) is part of a body of work done with Chondrus and other seaweeds that was done at the National Research Council of Canada where much has been learned about the metabolism of red seaweeds. Holding Chondrus crispus in conditions of high light and low nitrogen leads to an upward shift in the carbon/nitrogen ratio of the tissues and in a higher yield of kappa-carrageenan; perhaps also better quality in respect to gel strength (Neish & Shacklock, 1971). The photo above shows K. alvarezii plants from same-source propagules grown at different locations on the same farm at the same time. The white plants are from a low water flow area and have lost almost all pigmentation. (India, 2003) Various unpublished experiments on fertilisation have been carried out under farm conditions.
Under such circumstances there may be a huge number of water changes each day and usually very little effect of the fertilisation is Perhaps the same treatment of eucheuma seaplants may be seen even when the fertiliser is slowly disseminated from a porous worthwhile. It appears that this "ripening" phenomenon (also called container (Doty, unpublished). Mairh et al. (1986) and others have the "Neish effect" after Arthur C. Neish) may be related to had good growth and maintenance of stocks for a few years using macronutrient storage; in particular to nitrogen storage. This has not various formulations of artificial seawater and various micronutrient yet been demonstrated convincingly in eucheuma seaplants. mixtures. Fertilisation has never become widespread among eucheuma seaplant farmers; probably because of high fertiliser cost and high losses of fertiliser through diffusion when they apply it to sea cultures.
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4.H Micronutrients & Metabolites

Little is understood about the micronutrient requirements of eucheuma seaplants but they plants are clearly capable of assimilating and storing a wide range of elements including heavy metals and pesticide residues that must be monitored. During their studies of eucheuma seaplant growth in Hawaii Glenn and Doty (1990) noted that the micronutrient content of well waters ran from 50 to 100 times those of open coastal water and included a large amount of silica and iron. Some eucheuma seaplant species died and others thrived in waters whose silicate content is accepted as being indicative of strong igneous terrestrial influence (Lines 1 and 2, Table 4.F). Silicon per se probably has little direct influence on seaplants but it is associated with other elements that may effect them and silicate favors diatoms which can be pests in eucheuma seaplant cultures. Doty and others have published various reports inferring nutrient effects from the observed growth habits of plants on farms and in test plots. For example during early stages of farming in the Philippines floating cultures of Kappaphycus were repeatedly tried unsuccessfully and it was eventually concluded that something essential to growth was needed from materials generated near the sea bottom. This hypothesis was derived from results obtained in experiments in which plants were alternated between bottom and surface rafting (Doty, 1971b). In the northern Philippines Doty found that thalli grew poorly over the deep water but well where they were in contact with algal reef flats. For some years planting was done only on extensive algal reef flats. So far there has never been the combination of facilities for sampling and analysis that would enable one to learn what the bottom material contributes.
Despite the early negative Philippine experiences rafting has become widely accepted as a commercial cultivation method in most countries where eucheuma seaplants are farmed. It is almost universally used in some areas of Madura, Bali and Sulawesi in Indonesia and in some areas of the Southern Philippines where the sea bottom is unsuitable for "on/off-bottom" methods. The author has observed that raft-culture works best in areas where the seawater is very well mixed. This occurs in areas where strong tidal currents are found coursing through narrow channels in proximity to deep-water areas or near barrier reefs where seawater vigorously courses over the reef into farm areas (e.g.: in Bali; below) yet farm structures are protected from heavy wave action.
One can presume that Kappaphycus produces metabolic waste products that must be excreted from the thallus and diffused away from the plant. Kappaphycus spp. and Eucheuma spp. both exude a slight odour that intensifies on drying; especially in the case of Eucheuma. The odour is reminiscent of chlorine and may be associated with the brominated phenolic compounds that have been found in seaweeds. It has been speculated that such metabolites play a role in protecting plants from grazers.
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4.I Required Farm Site Characteristics

The basic elements of seaplant growth dictate that successful farm systems must have the following features: 1. Large surface area exposed to sunlight having optimum characteristics. 2. Effective, even water flow to and from all plants in the system. 3. Even dispersion of plants throughout farm sites. 4. Amenable to frequent cropping, cleaning and tending so weeds, pests, disease and fouling organisms cannot overrun seaplant cultures. 5. Rugged enough to withstand the substantial hydraulic forces of moving water and wind. 6. Located in places with environmental conditions as close to ideal as possible for the crops being grown. 7. Minimal fixed and variable production costs. 8. Protected from weather and sea conditions beyond farm habitats' structural limits. 9. Secure from human interference (e.g. pilferage, vandalism and accidental damage from boats).
Ultimately the only way to find out whether a given site supports vigorous plant growth is to plant test plots and expand where plants grow best. Only growing crops over several seasons confirms which locales are best. Site selection is critical to farm success. Site choice can lead to project failure or to success and competitive advantage. The critical factors necessary at a good site are: 1. Communities of people willing and able to become effective seaweed farmers. 2. Clean, nutrient-rich water at the right temperature. 3. Low probability of force majeur episodes due to natural or human causes. 4. Access to essential inputs, infrastructure and resources at attractive cost. 5. A stable, friendly climate for business, political and socioeconomic activities.
These factors have implications for the types of culture systems that can be used for seaplant cultivation. Examples: 1. Enclosures or suspension systems must expose the plants to both light and water flow so large volume-to-surface enclosures such as salmon cages cannot be used. 2. Plants must be separated and suspended by the use of physical structures (e.g. cages or ties on ropes) and/or applied force (e.g. air and paddle-wheel agitation). 3. Plants and animals can be combined in properly designed polyculture systems to their mutual benefit.
Aerial view of a eucheuma seaplant farm, Bohol, Philippines, ca. 1986
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4.J Farm Site Placement Strategies

The four major farm site placement options are illustrated in the figure 4.J below.
The four systems can be compared and contrasted as follows: Type A systems: Semi-enclosed or enclosed land-bound systems. Expensive to build and operate. Suitable sites often scarce or expensive. High tech in nature. Potentially enable a high degree of control over culture conditions. Systems often complex. Best suited to very high value crops for which control is worth the high costs. Type B systems: Tidal ponds such as used for prawns, milkfish and Gracilaria. Not used extensively for carrageenophytes due to water quality and environmental fluctuations. Sometimes embody worst characteristics of Type A and Type C systems. Type C systems: Most carrageenophytes are raised in Type C systems. Structures are placed in protected salt water bodies such as bays, channels and other inlets. Also used for Porphyra, Laminaria and other seaplants. Work well if placed where seasonal climatic shifts do not cause low growth, pest problems or death of the crops. Best Type C sites often near wave and current action.. Biggest problem is that suitable sites are scarce and prized for other uses (e.g. recreation, tourism, port facilities, other aquaculture uses). Type D systems: Open-water ocean systems. Essentially rugged and robust versions of Type C systems that can thrive in the exposed areas where weak or poorly designed gear cannot survive. Large available area and sites tend to be less effected by seasonality, grazing and some other problems of near shore operation.
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4. K Site Selection Tools and Data Products

By using the proper tools, data products and procedures one can select sites for farm development with minimal risk. It must always be kept in mind, however, that the installation and monitoring of test plots is essential before site potential can be established with certainty. The survey team must collect as much information as possible including tide tables or tide prediction programs (e.g. JTides); nautical charts of the area; historical weather records; and any oceanographic information that may be available. Information about local marine flora and fauna are invaluable in those rare instances where they are available. Basic survey tools include a good boat, the items illustrated below, radios or cell phones and instruments used for monitoring the environment. A good set of nautical charts and tide tables that cover your survey area will save much time and effort. in spotting likely farm sites.
All surveys must include getting in the water and "ground truthing" to see what is there. Here the author is looking for sites near Semporna, Sabah where farms later developed commercially. (1977 Jack Fisher photo) Nothing beats aerial surveys for overviews of farm sites and monitoring of plantings. Photo top right shows the author about to take off in Piper Cherokee RPC1252 for a survey and photo session over seaplant farms in the central Philippines (photo lower, right; ca. 1984). Most light, microlight or ultralight aircraft can be used for this function.
Mask and fins
rugged GPS
water proof Pelican case
Marine binoculars
rugged computer
manufacturers' photos
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5. A
Agronomy Overview
Elements Requiring Management People
The fundamental basis of seaplant agronomy is that a farmer works with other people to create habitats suitable for the desired crops within natural environments. The farmer uses agronomy methods to monitor the status and behaviour of these fundamental elements and then, to the best of his or her ability, the farmer manages and controls the farm system to profitably produce a saleable crop.
The social environment in which the seaplant enterprise takes place, especially stakeholders such as , employees, suppliers, buyers, government agents etc.
Environment Interactions with the physical environments of seaplant farms especially minimisation of adverse impacts. Habitat Crop Development, construction and operation of the physical structures that comprise the seaplant farm. The population of eucheuma seaplants that comprise the farm populations and yield the crop.
Monitoring functions People

Semporna, Sabah, Malaysia, 1996
Maintaining useful interaction with stakeholders.
Environment Measurement, recording and data handling for meteorological and oceanographic parameters. Habitat Crop Oversight and inspection leading to preventative maintenance of farm structures and equipment. Crop logging.
Elements and functions of this process are shown in Fig. 5.A below and are further described in Table 5.A opposite.
Control functions People Standard personnel management and business management for small-medium farm enterprises.
Environment Pro-active minimisation of negative environmental impacts and maximisation of positive impacts. Habitat Crop Operation of farm structures and equipment through manually applied operational protocols and automated control systems. Planting, maintenance and harvesting and post-harvest treatment of the crop.
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5. B
Some Background
The commercial success of eucheuma seaplant farming is based on the fact that these plants produce vegetative thalli large enough to be economically planted as propagules (cuttings) and harvested individually.
The attachment methods initially used for fastening cuttings to lines were the "tie-tie" method and containment in bags.
Since 2000 a "loop" system has been introduced in Madura, Indonesia. The "loop" system originated by Mr. Made This insight surfaced during experimental farming conducted in the Philippines during the late 1960s and the early 1970s. The history of Simbik of Bali (photo right) reduces planting labour, eases the recycling of exactly who discovered what and how strains such as "tambalang" planting materials and eliminates most came to be is rather muddled but it seems to have arisen from raffia or string from the crop. activities jointly and variously undertaken by individual farmers working in concert with Doty's University of Hawaii teams; the Philippine Bureau of Fisheries and Aquatic Resources (BFAR); various Philippine universities; and Marine Colloids Philippines Inc. Bag methods protect crops but they are expensive so their use tends (now owned by FMC Biopolymer). to be confined to propagule production. The major habitat During the development of farming in the Philippines early methods configurations that developed for commercial use have been the "floating" and "off-bottom" systems. included attaching cuttings to stones but this method soon gave way to tying cuttings on nets. Nets quickly gave to monolines because these are more convenient to handle. This work established farming by methods that involved thalli held rigidly held in place within porous bags and/or tied directly to lines held taut between supporting structures. Such methods are fundamentally different than the tumble culture popularly used to produce pond cultivated Chondrus and other smaller seaplants. When "attach-to-string" methods are used labour comprises most of the cost of crop production. "Attach-to-string" farming methods facilitate experimentation and cultivar screening. Individual test thalli can be labeled, removed, weighed and replaced at intervals. The growth rates of individual thalli can then be easily calculated from successive weights This approach greatly facilitates comparisons between farming strategies and crop varieties so farmers are able to innovate and expand farms rapidly. Floating methods employ rafts (usually made of bamboo) or systems of floats (most often empty plastic beverage bottles) to suspend lines near the sea surface. Off-bottom methods utilise stakes driven into the sea floor to suspend lines above the sea floor. General advantages of floating systems include: 1. Grazing by bottom associated animals is minimised or eliminated because the plants are raised out of reach of the benthic grazers. 2. Plants near to the surface of the water column are often exposed to more adequate water movement (e.g. wave chop). 3. Floating cultures can be tended during any tide level whereas work on off-bottom cultures is limited by tidal cycles. 4. Floating cultures are not restricted to shallow waters. A wide variety of line arrays can be seen in commercial farms but minimum line spacing is seldom less than 15 cm. Sizes of farms and rafts vary widely. Efficacy of habitat construction and technique is sometimes dictated by environmental factors but in other cases it seems to be a matter of farmers' experience or preference.
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5. C
Types of Habitat System
The most obvious physical manifestation of seaplant farms is their habitat structures.
Seaplant farm habitat characteristics; list and descriptions: Substrate or enclosure type
Propagules attached directly to stones or coral fragments using string, elastic bands or netting. Generally for temporary holding pf propagules Fishing line or twine; usually 3-8 mm diameter Nets bags or tubes made from fish net or agricultural netting Plastic bag or tubes with or without perforations Ponds or raceways built on land; water fed by tide or pumps
These incur most capital and operating costs so it is normal for farm coral/stones systems to be classified according to their most obvious physical characteristics such as "off-bottom" or "bamboo raft" or "longline", pen/cage/sack etc. Ask and Azanza (2002, Table 3) have summarised descriptions of monoline several such farm systems. net bag/tube The table opposite describes methods considered in the present film bag/tube monograph. pond/raceway on off bottom mid water surface
Horizontal net culture, Kappaphycus alvarezii, Sulu Sea, 1977
Position or location
Propagules placed on sea floor Propagules placed just above sea floor Propagules placed in water column Propagules floating at or near sea surface Farm systems placed in ponds on land; intertidal or above tides
on land
Orientation The fact is that the diversity of farm habitat systems has gotten so Substrate horizontal to sea floor/surface diverse that they can no longer be meaningfully described or in terms horizontal of just one or a few of their characteristics. Habitat systems comprise perpendicular Substrate vertical / perpendicular to sea floor/surface combinations of five features such as: Propagules unattached; mixed by physical agitation mixed 1. the type of substrate of enclosure that serves as a physical Fixation method matrix for holding crops within farm boundaries. stakes/rocks Support substrate or enclosures on or near sea floor 2. 3. 4. 5. the position or location of the substrate or enclosure relative to the sea floor and the sea surface. whether the substrate or enclosure is oriented horizontal or perpendicular to the sea surface (or has no fixed orientation). the method by which the substrate or enclosure is fixed, suspended or held in place within the farm area. the method by which propagules are attached to or suspended within the substrate or enclosure. floats+anchors Support substrate or enclosures in water column or at/near sea surface rafts terrestrial none enclosure tie loop
Substrate or enclosures suspended from bamboo or plastic-tube rafts Farm systems placed in ponds on land; intertidal or above tides
Attachment method
Propagules unattached; mixed by physical agitation or free floating Propagules individually unattached but contained in enclosure Propagules individually tied to substrate using knots Propagules individually attached to substrate using loops
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5. D
Substrate coral/stones
Broadcasting and Tie-on-stones Systems

Orientation horizontal Fixation stakes/rocks Attachment none or tie
Position on bottom
Wild Kappaphycus plants growing naturally near Semporna, Sabah, Malaysia, 1977. Right, centre: Wild Kappaphycus with the flattened appearance of the K. cottonii type. Growing in typical habitat among corals, sea grasses and other seaplants
Eucheuma seaplants were harvested from wild stocks in China, Indonesia, the Philippines, Malaysia, Tanzania and Indonesia commencing in the mid 20th century. Total Quantities never exceeded about 1,000 tons per annum and the harvests were generally of mixed species. Early cultivation efforts involved broadcasting of cuttings over the sea floor; attachment to stones or coral fragments using string or elastic bands; or sandwiching cuttings to the sea floor using nets to make a "lawn". Betaphycus gelatinae cultivated on stones and coral, Hainan, China 1988
Photos above and below by Jack Fisher, 1977
Right: A wild "On-bottom" methods caused localised population increases but were Kappaphycus plant of the K. laborious, disrupted the benthos and allowed little crop control. striatum type. Grazing by It is selection sea urchins from wild and other plants of this herbivores appearance was a major that led to problem "oncurrent bottom" (see commercial photo right). strains of Kappaphycus.
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5. E
Substrate
Nets and Monolines in "Off-bottom" Systems

Orientation Fixation Attachment
"Off-bottom" monoline systems vie with floating systems as the preferred habitat structures at most farm locations. monoline or net off bottom horizontal stakes/rocks tie or loop This sort of scene State-of-the-art during the early 1970s in the Philippines and Sabah has become a was the use of large-mesh nylon monoline nets with propagules local low-tide tied at net junctions about 40 cm apart. feature of beaches in The nets were generally supported just above the sea floor by tethers that secured them to coral heads, rocks or mangrove-wood eucheuma seaplant farm stakes driven into the sea floor; generally over the types seagrass areas. beds that serve as natural eucheuma seaplant habitat. Bamboo rafts were also used to a limited extent; usually on test plots.
Position
K. alvarezii growing on netting on a test plot operated by Vic Alvarez in Sabah (1977). The "tambalang" variety probably originated near here.
above: Tanzania, 1991
Lines may be suspended above the sea floor using stones (above left; India; 2003) or stakes made from concrete, steel or rot resistant wood (above right; Environmental impact Tanzania; 1991). Stakes are Mangrove wood is commonly used as stake material but cutting this normally driven into a soft is banned in many places for sound environment protection limestone seafloor; often with prereasons. drilled holes (Bali, right; ca. 1989). The system of using large-mesh nets fell into disuse when it was found that plants could be spaced as little as 15-20 cm. apart. Nets of this mesh are laborious to make, impede walking through farms and offer no advantage over monolines.
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5. F
Enclosure
Pen, Cage, Sack Bag and Tube Systems

Orientation Fixation Attachment
Net sacks (right) or pens are commonly pen/cage/sack on/off bottom horizontal stakes/rocks enclosure used for holding net bag/tube mid water perpendicular rafts or floats tie or loop propagules after film bag/tube surface mixed terrestrial harvesting or before planting. Sometimes Since the beginnings of eucheuma seaplant cultivation a plants are grown inside bewildering variety of systems have been tried. These have cages (below, placed loose or tied plants within pens, sack, cages, bags or tubes Philippines ca. 1990). made from netting or perforated plastic.
Position
The use of such systems has been tried with several species of seaplant. Their use is sometimes motivated by attempts to reduce the labour of attaching or confining plants; especially with small species (photo below). Other motives include protection from grazers and prevention of plant losses through breakage. Chondrus crispus growing in a bag in quarantine. Mexico; 1996. Although "enclosure" methods are commonly used for temporary holding of live propagules they have never gained popularity for crop grow-out. For various reasons their efficacy seldom offsets the high capital and operating costs that such systems have relative to "attach-to-string" methods. The jury is still out on "enclosure cultivation. Variants of such systems may prove to be cost-effective as technology and agronomy protocols advance.
Eucheuma seaplants have been experimentally cultured using several types of tubing including the "chorizo" style shown here in use with Chondrus crispus. Growing fronds may protrude from bags or remain inside where loose plants can suffer tip abrasion (above, right).
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5. G
Substrate monoline net/film bag or tube
Float + Anchor Long-line and Short-line Systems

Position mid water surface Orientation horizontal perpendicular Fixation floats + anchors Attachment tie or loop enclosure
The "long-lines" illustrated below work well where large areas of water deeper than about two metres are available for farms.
Floating short-line or long-line systems have found favour especially where eucheuma seaplants are grown in deep water. This method can be among the most environment-friendly means of growing eucheuma seaplants. Farms can be placed in deep water (e.g. 3 metres or more); farms can be planted over muddy or sandy bottom where plant fragments do not attach; and boats are used to minimise sea floor damage.
Lines can be fastened closely together if need be (photo below).
Farmer tending lines near Semporna, Sabah; 1996
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5. H
Substrate monoline net/film bag or tube
Raft Systems
Position surface Orientation horizontal Fixation floats + anchors Attachment tie or loop enclosure
One advantage of raft systems is that units can be brought to shoreside work areas and the need for work on the water is minimised. This saves on costs and reduces safety hazards.
Raft systems have found favour where eucheuma seaplants are grown in areas with plentiful, large bamboo and/or sea floors that are inappropriate and/or too deep for on/off-bottom systems. This method can be also be environment-friendly. As with other floating systems farms can be placed in any depth of water; farms can be planted over bottom where plant fragments do not attach; and boats can be used to minimise sea floor damage.
These 10 x 10 m. rafts at Madura, Indonesia (above) are part of a 30 km coastline that is virtually continuous eucheuma seaplant farm and supports at least 1,000 farm families. The 2.5 x 5 m. rafts below are at Gunung Payong, Bali. Bamboo is normally used but plastic piping can also be utilised. Made Simbik inspects his farm at Geger, Bali, Indonesia (ca. 1989). This area was the FMC "Seaweed University" at that time.
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5. I
Enclosure pond/raceway net/film bag/tube
Terrestrial Pond and Raceway Systems

Position on land Orientation mixed/agitated Fixation terrestrial Attachment none enclosure
Propagules may be unattached and agitated; they may be attached to substrate within tanks or raceways and/or they may be placed within sub-enclosures such as nets and tubes. As in the case of net and tube systems, some variants of pond or raceway systems may prove to be cost effective for cultivation of eucheuma seaplants as technology and agronomy protocols advance or as commercial conditions change. One common means for maintaining unattached propagules in suspension is a stream of air bubbles from a blower (right).
Experimental Chondrus crispus systems; Marine Colloids Ltd./NRCC; 1976
Terrestrial pond and raceway systems have been used for experimental cultivation of eucheuma seaplants but have not been employed for large-scale cultivation. Terrestrial pond and raceway systems permit a high degree of control over culture conditions but this is achieved at high capitaland operating-cost.
Tank and raceway systems are useful for experimental work, however, and can also be used in seedstock election programs (above, Bali, ca. 1991). From time to time attempts have been made to integrate eucheuma seaplants into land-based polyculture systems that include fish, crustacea and molluscs (e.g. below) but this approach has not been widely adopted.
Another common means for maintaining propagules in suspension is paddle wheels such as this one in Nova Scotia, Canada (right). Agitation involves substantial energy consumption. That tends to restrict these systems to high-value crops.
Experimental polyculture systems that incorporated eucheuma seaplants (Kahuku, Hawaii ca. 1981. Jack Fisher shown working above.)
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5. J Positions and Orientations of Habitat Systems

Substrate/enclosure Position coral/stones pen/cage/sack monoline net bag/tube film bag/tube pond/raceway on bottom off bottom mid water surface terrestrial Orientation horizontal perpendicular mixed/agitated Fixation stakes/rocks Attachment none
floats + anchors enclosure rafts terrestrial tie loop
The orientation of habitat systems is generally horizontal to the sea surface and the sea floor but in some locations conditions permit a perpendicular orientation.
The range of feasible positions and orientations for eucheuma seaplant habitat-systems is shown in Figure 5.J below
The Bali farm illustrated above (ca. 1989) illustrates the horizontal orientation typical of most farms. Vertical systems may be "topdown" with lines hanging from the surface or, more commonly, bottom up with lines floating toward the surface as they are fixed to the sea floor and suspended by floats (e.g. Philippine farm at right; ca. 1990). The choice of position and orientation is governed by a host of local factors including space constraints, water motion, water depth, ease of handling and cost of materials. As a general rule horizontal systems with lines held rigidly in place are preferred because such systems minimise line tangling and are generally easiest to handle during planting and harvesting. The "agitated" or "mixed" type of orientation is illustrated in the monograph section that describes pond and raceway systems.
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5. K Attachment Systems
Substrate monoline Position on/off bottom mid water to surface Orientation horizontal perpendicular Fixation stakes/rocks rafts, floats + anchors Attachment tie or loop
A "loop" method that seems to have originated in Bali or Madura has several advantages relative to the practise of tying propagules using slip-knots or other types of knot. First, the labour involved in attaching propagules is much reduced. Second, harvesting is accomplished by simply drawing the planted line through a hole in a piece of wood (right) to cut plants without losing "loops" and mixing string in the crop. Finally, string with loops can typically be recycled for several cycles. Loops of about 2 cm are secured by one end through the line plaits at intervals of about 15-20 cm. The other end of the loops is left loose. To plant push line through the loop; insert propagule; pull line snug.
Overwhelmingly cultivation of eucheuma seaplants involves attachment of individual propagules ("cuttings" or "Seedlings") to pieces of string (lines). This is the most labourious aspect of eucheuma seaplant farming.
On average 3-4 hours/day are spent tying propagules to lines. In recent years the use of monofilament lines (usually nylon) has given way to the use of plaited lines (usually polypropylene). Plaited line tends to be cheaper and less prone to tangle but it is less durable than nylon monoline and tends to disintegrate rapidly if exposed directly to sunlight. Line is generally sold by weight and the cost per unit of length is exponentially related to line diameter so farmers tend to use the smallest line that they can. Typical current costs for polypropylene line are shown in Figure 5.K right (US dollars/km).
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5. L Seasonality - Periodic Variations in Growth

Variation in growth has been observed for Kappaphycus both within and between annual periods. Productivity variations that appear more or less as regular annual cycles are commonly referred to as "seasonality". Growth differences between years tend to be associated with macro-climatic phenomena such as el Nio or la Nia. Generalised variations in farm productivity were observed early in the development of eucheuma seaplant farming (Doty, 1987; Table 5.L below) but these phenomena are still poorly understood. Some seasonality appears in most commercial eucheuma seaplant farming areas and direct effects on the crop seem to be related to meteorological and oceanographic phenomena such as wind direction, water movement, nutrient availability and temperature changes. Secondary phenomena include seasonal grazing activity and pest, weed or disease problems associated with seasonal phenomena. 1976-77 Mean Specific Growth Rates (% day-1) (Doty, 1987) Spring Summer Fall Winter Water-in (avg.) Water-out (avg.) K. alvarezii 5.25 6.23 4.41 4.15 5.05 2.07 K. striatum 3.95 3.33 2.98 2.98 3.51 1.43 E. denticulatum 3.51 4.05 3.52 3.35 3.61 2.29
When farm areas exhibit strong seasonality effects there is little that can be done other than to relocate the farm. When and how to do this is determined my the effective use of environmental monitoring and crop-logging programs (whether intuitive or formalised) and the development of effective controls or remedial actions. In general farmers have three choices if farming becomes becomes seasonally uneconomic at particular sites. They are: 1. Harvest and sell. 2. Move the farm to another site. 3. Transfer thalli into a relatively deep hollow that has a low but sustaining light intensity; then wait for a favourable seasonal change. The decision as to which course to take depends on current market conditions and the availability of suitable alternate planting sites. With respect to long-term farm productivity variations it seems that little is understood. Attribution of effects to el Nio or la Nia attach a name to such phenomena but contribute little by way of explanation especially where there is weak correlation to begin with. For example during 1998-1999 poorer-than-previously cottonii growth was reported to the author by various informants in sites as widespread as the Sulu Sea and Zanzibar, yet the productivity of spinosum (E. denticulatum) continued undiminished. In some areas the commercial cultivation of cottonii ceased and spinosum replaced it where market conditions permitted.
In most tropical regions of the Indian and Pacific Oceans there are seasonal shifts in prevailing wind direction and general weather conditions generally referred to as "monsoons". In some farm areas the monsoons cause acceptable variations in farm productivity but in others there may be changes that make the farming of certain sites uneconomic during some seasons.
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5. M
Starting farms - the Crucial Role of Test Plots
As farm development projects go from initial appraisals to rapid expansion they pass through a highly critical period that is fraught with risk.
Eucheuma seaplant health is not difficult to read. Growth, colour, cleanliness, shape and gum quality usually tell all.
Seaweed growth results from a complex combination of effects but overall plant health can be read by using readily detectable parameters such as growth, colour, cleanliness, shape and gum quality. For example nitrogen is often a limiting nutrient in seawater and some combinations of environmental factors can cause the rate of photosynthesis to exceed the rate of nitrogen assimilation. As plants metabolise nitrogen stored in their pigments the loss or gain in There are no experts on commercial seaweed farming at new sites. colour density over time is a good indicator of plants' nitrogen status. Initially the best one can do is to interpret plant behaviour because: When developing new areas install plenty of test plots Project participants are early in the learning curve; areas with unknown suitability are being developed; seasonality is poorly understood; complex community relations issues are sure to arise at many sites; both sincere and opportunistic environmentalist groups are watching with a critical eye; and logistic strategies are still undergoing refinement. 1. Like most new parents, you do not know what you are doing. As new areas are developed farming is usually being undertaken at an unprecedented intensity and in poorly known geographical locations. 2. Inevitably plant growth and farm dynamics will vary seasonally between sites and among agronomy protocols. 3. There is general body of knowledge as to how seaweed crops grow but not much specific knowledge is in place. Seaplant scientists can usually only advance hypotheses or theories that require further testing. Inevitably the farmability of sites will vary among locations, between seaweed varieties, through seasons and between different crop years. This applies to any crop. Clearly it is better to detect problems on a small scale rather having potentially high-profile failures at the level of multi-hectare sites. Test plots have ongoing value in the planning of farming tactics. Proper crop-logging of plants on test plots will give guidance as to what corrective actions may provide solutions as conditions shift. Expand farms using small plots at many places; clearly identify these as test sites; avoid perceived failures
The sight of dozens of people in the sea and on the seashore doing 4. Typically one must adopt and empirical, pragmatic farming can make a powerful impression. This can be a two-edged approach to commercial farm development this is the way that sword. The inevitable failure of some sites has the potential to be successful farmers operate throughout the world. high-profile and may lead to adverse spin-off. Starting small at many sites rather than going large at fewer sites is initially awkward from a Solutions to these problems include the following: management point of view but reaps substantial rewards in the long 1. Develop a system of test plots and crop logging. run. This approach quickly reveals the most cost effective sites. 2. Tend crops diligently. Environmental and business impacts 3. Develop crop-sense by being sensitive to what the crop is telling you so variances can be corrected quickly. 4. Zero in on the best sites using the information generated from field trials. Remember - dont rush "mother nature". Expanding too far, too fast with limited site experience can lead to a world of grief.
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5. N Seaplant Agronomy and Crop-Logging Basics

Essential basic concepts Sustainable eucheuma seaplant farming produces bountiful crops on a sustainable basis with minimal disruption to surrounding environments. There are two "rules of thumb" that comprise the most basic concepts of this approach: 1. Go with the flow of environmental conditions and minimise environmental disturbances. Seaplant farming is among the arts and sciences that conform to natural conditions - it is not a set of methods for "bending nature to your will". 2. Observe and learn from what the crop and associated organisms are doing. Learn how to interpret the "signals" that they are sending out and plan actions accordingly. Successful eucheuma seaplant farming requires that you go with the flow of seasonal events by taking these steps: 1. Selecting several locations that permit seasonal shifts of farming effort among locations - a defining characteristic of sea farming. 2. Designing farm equipment and agronomy protocols that facilitate flexibility in farm expansion, contraction and re-location. 3. Implementing a crop-logging system that enables timely tracking of seasonal changes and indicates appropriate action. Crop-logging is a process of recording environmental variables and crop condition in a structured and methodical manner. Results must be saved in databases and results must be circulated to technical and management personnel. Good statistics should be kept for farm inputs and outputs. This is macro-crop-logging information that can give valuable insights crop logging done at the micro level.
As experience is gained with seaplant crops it becomes rather easy to "read" what sort of condition they are in. Even a beginner can probably rank the following plants in order of their well-being:
the good
the bad
and the ugly
A good crop log database comprises valuable equity for any seaplant farming business. Proper development and use of this information preserves experience and knowledge through successive generations of management and staff. Your hard-earned expertise does not simply "walk out the door" when people move on to other things or get pirated by your competion and new recruits do not have to waste valuable resources reinventing the wheel. Successful crop-logging involves vigilance and methodical recording of data using three types of monitoring program: 1. Keeping accurate records of planting, harvesting and growth rates (e.g. through use of test-lines and test-plots). 2. Measurement of oceanographic and meteorological parameters. 3. Monitoring of crop condition. Ideally sites should be monitored on a weekly basis. The crop-logging program should be augmented by use of properly stored and indexed digital photographs.
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monograph index Page 50

XYZ
5. O Crop Logging Parameters and Actions

AYZ Pink
next
Crop logging need not be complicated or technical. Main parameters to monitor are shown in the paradigm below.
Purple
GRAZING + BLEACHING + W.E.E.D. - Early symptoms of bleaching; noticeable grazing + W.E.E.D. AYC Cyan GRAZING + BLEACHING Plants show early symptoms of bleaching and show signs of noticeable grazing. ABZ Tan W.E.E.D. + BLEACHING Early symptoms of bleaching; Weeds, Epiphytes, Epizoa or Disease. ABC Yellow BLEACHING - Plants may appear to be healthy but have a very pale colour. Growth is slowing or becoming negative. Essential nutrients are depleted. Ice-ice symptoms likely.
GRAZING + W.E.E.D. Plants healthy; good colour many grazers + W.E.E.D XBZ Blue GRAZING - Plants healthy; good colour; noticeable grazing. Significant grazing means significant losses; severe grazing means grazers harvesting most of crop. XYC Brown WEEDS, EPIPHYTES, EPIZOA OR DISEASE Plants have good colour but if W.E.E.D. become significant farm economics suffer. At severe levels it is time to crop out and wait. XBC Gold GOOD AS GOLD - Plants are healthy, with good colour and no more than trace amounts of grazers or W.E.E.D.
Action options available in response to observed condition and behaviour are as follows:
GREEN = MAXIMISE LIME = MAINTAIN AMBER = BE VIGILANT Tend the crop industriously and take advantage of the good crop yields that come with Condition Gold. Plant vigorously. The ability to expand plantings during Condition Gold will determine maximum attainable farm yields. Tend the crop industriously and maintain vigilance to ensure that "condition yellow" is not emerging. Plant vigorously as long as signs of problems are not worsening at a noticeable rate. Be ready to harvest as soon as there are signs of impending or rapidly developing trouble. Maintain frequent vigilance to see if worsening conditions are progressive and worsening. Re-seed with better propagules or adjust agronomy protocols if possible. The crop is still doing OK but there are losses to grazers or noticeable W.E.E.D. problems that add to labour cost (e.g. having to pick out weeds) and/or reduce crop quality (e.g. epiphyte/epizoa contamination of crop).
ORANGE = MOVE If possible move crop to better sites if condition is tending toward condition 3 and/or growth decreasing. Conditions can rapidly move toward an Action Red situation where significant crop losses may occur. RED = BAIL OUT Move crop immediately to sites with better conditions or crop out farm. If conditions are such that Action Red is required that means that significant crop losses are occurring.
A printable table that matches actions to crop indices is shown as Table 5.P.
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5. P Index & Actions Chart for Kappaphycus and Other Eucheuma Seaplants
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5. Q Bleaching as a Crop Logging Index

Severe bleaching is BLEACHING - Plants may appear to be healthy but have a very pale occurring with colour. Growth is slowing or becoming negative. Essential nutrients are depleted. Ice-ice symptoms are likely to develop if the condition the plant at right. The plant persists. is clean and intact but its nutrients are These plants are already from the same depleted. The place and from the plant is not same stock but at growing and will different seasons. soon die. The plant at top is Moving a plant in "condition gold". such as this to a nutrient rich A trend toward area will bleaching is generally result indicated in the in full recovery. plant below. Colour density is low but the plant is still healthy and "False growing well on the bleaching" is borderline of occurring in Condition Gold. the frond at This plant is in an lower right. area that later The fronds developed the cortical layer severe bleaching has actually shown at upper been stripped right. off by grazers. ABC Yellow ABZ Tan AYC Cyan AYZ Pink
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5. R Types of Grazer Damage

XBZ Blue XYZ Purple AYZ Pink AYC Cyan Tip-nipping occurs when growing tips are bitten off. In the plant at right all tips have been nipped but new ones are growing back. Tip nipping is commonly seen and is often attributed to fish such as rabbitfish and juvenile surgeonfish or parrotfish. Total damage can occur as in the plant shown tight. This plant seems to have incurred all of the types of grazer damage described here except total loss. GRAZING - Plants healthy; good colour; noticeable grazing. Significant grazing means significant losses; severe grazing means that grazers are harvesting most of your crop. Gouging is the sort of damage where small chunks of pigmented tissue are removed as on the thallus shown right. This pattern seems typical of snails and perhaps of small sea urchins. Planing is the sort of damage where the side of a branch is flattened by removal of tissue as if by a plane. This pattern seems typical of the larger sea urchins.
Stripping occurs when gouging or planing is so severe as to cause complete removal of the plants' pigmented cortical layers as at right.
Plants just gone as if chomped off in one bite? This can be done by green turtles (Chelonia midas). These animals can be kept out of farms by sturdy fences but avoid planting where they feed if you can.
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5. S Examples of Grazers
XBZ
XYZ AYZ AYC GRAZING - Plants healthy; good colour; noticeable grazing. Significant grazing means significant losses; severe Blue Purple Pink Cyan grazing means that grazers are harvesting most of crop. Grazing by herbivorous marine animals has been a problem for eucheuma seaplant farmers since the beginning of agronomy development (Doty, 1973; Parker, 1974; Doty and Alvarez, 1981) but there have been few published studies of this topic. Grazer sizes range from the very small (e.g. apple snail, near right) to the fairly large (e.g. green turtle, Chelonia midas, far right). The presence of grazers is usually due to the placement of farm habitats in or near to seagrass beds and other areas where marine herbivores have endemic populations. In other cases herbivores may be introduced through the transport of propagules among farm locations. In still other cases the herbivores come to farms when schools of fish pass through a farms during certain seasons or life-cycle stages. The sea urchin species Diadema setosum and Tripneustes gratilla are commonly found in the vicinity of eucheuma seaplant farms. They appear to be "planers", "gougers" and "strippers". If farm habitats are placed on or near the sea floor for several months or years it appears that localised "population explosions" of grazers can be stimulated (author; pers. obs.). Diadema setosum Tripneustes gratilla parrot fish; Cetoscarus bicolor rabbit fish; Siganus javus surgeon fish; acanthurus_mata
Russell (1983) and Uy et al. (1998) lent formal confirmation to common observations of farmers when they observed that fish selectively graze smaller branches of eucheuma seaplants. Juvenile parrotfish (scarids) and surgeonfish (acanthurids) have been observed to consume as much as 50-80% of eucheuma seaplant populations at 0.5 and 2.0 m depth (Russell, 1983) and rabbitfish (siganids) are commonly seen to be feeding on eucheuma seaplants. Most fish seem to be "tip nippers". Other herbivorous fish families including puffers fish are known or suspected to be eucheuma grazers and a wide variety of invertebrates including holothurians and crustacea are commonly seen in the vicinity of eucheuma seaplant farms. The author has received numerous reports of star fish and sea cucumbers eating seaplant crops. Herbivory on eucheuma seaplants is a subject with abundant scope for useful study. Environmental imact If you do not place your farm in the midst of grazer habitat they are far less likely to bother you.
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5. T Control and Prevention of Grazing

XBZ Blue XYZ Purple AYZ Pink AYC Cyan
"Brute force" to combat grazers runs counter to the "go with the flow" approach that has made for economic success in the organic Attempts to "engineer" ways out of grazing problems have generally production of eucheuma seaplants. During more than three decades proven to be expensive and futile. of dealing with grazer problems in seaplant cultures the author has Environmental impact reached the following conclusions as to why several "red" pest controls fail: Grazing can be stopped in a variety of ways but few methods are both environmentally acceptable and economically feasible. The best 1. POISONS - Pesticides can be effective in ponds, raceways or other "green" ways to deal with grazers are to: enclosures but many poisons may be needed to combat the wide spectrum of vertebrates and invertebrates found eating seaplants. Also, like fertilisers, 1. AVOID THEM - Place farm habitats in locations where endemic grazer they diffuse rapidly in the sea so in situ application results in most being populations are not abundant; for example by placing floating habitats in wasted and drifting off to contaminate surrounding environments. water several metres deep over muddy or sandy bottom. 2. CHEMICAL DETERRENTS - May work briefly but diffuse and contaminate 2. SWAMP THEM OUT - Build eucheuma seaplant populations to a "critical surrounding environments. Herbivores tend to habituate to the presence of mass" where any grazing pressure is trivial relative to the total biomass and chemicals and other deterrents. production; then accept some losses. This is a widest-spread method of dealing with grazers. 3. AUDITORY, VISUAL, ELECTRICAL AND OTHER DETERRENTS - Fish are not as stupid as most people seem to think - they can habituate and 3. BLOCK THEM - Use barrier nets or enclosures on a selective basis during adapt rapidly to a wide variety of deterrents, especially "scarecrow-style" seedstock production or to prevent the entrance of large grazers such as devices. Some devices, such as palm fronds, may act as deterrents at first turtles. then as attractants later. Many invertebrate grazers probably also habituate. 4. EVADE THEM - Crop back and wait out seasonal grazing periods if they last Besides being potentially ineffective, deterrent devices may disrupt for only a few weeks (e.g. in Bali, Indonesia; pers. obs.). populations in surrounding environments. 5. CATCH THEM - follow the 4. BARRIERS & TRAPS - Can keep herbivores out or pen them in depending example of these seaplant on how they are used. Without a doubt the selective use of barriers and farmers in Sabah. Cottonii is traps can have a role in seaplant farming (see "block them" opposite) drying on the platform (top especially for highly seasonal grazer phenomena, for very large grazers or of photo), the kids are for protecting small plots of high value plants such as a cultivar bank. For waiting to eat and the the most part they are too expensive too install and maintain as a mothers are preparing component of crop production systems. freshly caught rabbit fish for Environmental impact lunch. Many marine herbivores are good to eat. Fortunately there are vast tracts of inshore areas where seaplant Some have aquaculture farming has been successful without resort to drastic grazer control potential (e.g. abalone; measures. Find them and use them. rabbitfish).
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5. U Diseases and Malnutrition

XYC Brown Communicable diseases leading to epidemics are not known to have occurred among eucheuma seaplants but maladies attributed to malnutrition are commonplace as a seasonal event in many areas. Remedial action is usually to "bail out". The malady known as "ice-ice" was identified as a problem early in industry development (Doty and Alvarez, 1975; Uyenco et al., 1981). The term was coined by Filipino farmers to describe the dying tissue devoid of pigment that causes branches to break off (photo right). Doty (1978b) identified stress as the major factor promoting ice-ice and drew a correlation between its occurrence and that of epiphytes. The progress of the ice-ice phenomenon is generally as follows: 1. Formerly dark-coloured, clean, vigorously growing plants lose pigmentation while otherwise remaining healthy in appearance and growth rate. 2. In a matter of weeks; sometimes even in a matter of days; the loss of pigmentation may become severe and growth rate becomes very low or negative. At this stage if the plants are moved to "better water" they may exhibit full recovery. 3. New tips, if they occur at all, tend to be spindly and lacking in vigour. In this case the plant has acquired an overall appearance that is reminiscent of a centipede. There were many tips but growth was negligible.
4. Areas devoid of pigment appear at intervals on plant thalli (below left); they weaken and eventually the tissue atrophies, thus causing plant breakage (below, right).
Uyenco et al. (1981) noted high populations of bacteria found on tissue with ice-ice but concluded that they were secondary to the problem. Doty (1987) noted that the occurrence of ice-ice was seasonal and was correlated with changes in the monsoon (Doty and Alvarez, 1975). Largo et al. (1995a, b) showed that certain bacteria appeared to be capable of inducing "ice-ice in stressed propagules and noted that that several abiotic factors could generate symptoms. Light intensity of less than 50 micromol photon m2 s; salinity of less than 20 ppt.; and high temperature (up to 35oC) induced "ice-ice" in K alvarezii planted in sub-tropical waters in southern Japan (op.cit.). Eucheuma seaplant metabolites may also play a role in ice-ice formation. Mtolera et al. (1996) observed that E. denticulatum produced volatile halocarbons (VHC) in high-light and low-C02 environments . Pedersen et al. (1996) demonstrated that high VHC levels produced diseases in Gracilaria cornea and a similar phenomenon may occur with eucheuma seaplants, especially in low-water-flow; high-unmixedboundary-layer conditions.
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5. V
XYC
Weeds, Epiphytes and Epizoa

ABZ Tan XYZ Purple
Brown
W.E.E.D. - If weeds, epiphytes, epizoa or disease become prominent you should crop out the farm and move it and/or replace propagules with healthy material. Weeds are algae that grow in a mix with the desired crop. If weeds are not too they can be picked out of the crop before drying (below).
Settling of loose silt and/or microalgae is common in areas with murky and/or still water. Combat this by shaking plants (daily if possible) during times of tidal flow.
Various Ulva spp. (upper, left) are common seasonal weeds. Severe weed infestations (upper, Seasonal weeds were noted as a serious problem from the beginnings right) may smother crops. of eucheuma seaplant cultivation (Parker, 1974) but the problem is Animals such as bryozoans may still not well studied. Fletcher (1995) provided a review on the also grow on the crop (right); impacts of pest weeds on Gracilaria cultivation that is relevant to the especially on old tissue. These, situation with eucheuma seaplants. animals diatoms and other small plants or animals can form a Three types of pest weed can be recognised: "scum" on the crop that 1. Macroalgae such as Ulva spp., Enteromorpha spp., Cladophora diminishes growth and devalues spp. and several other genera that drift or settle; then tangle or the crop. attach to farm habitat structures or to the crop. Usually acutely Current protocols for managing non-epiphytic weeds call for watching seasonal with duration of a few weeks or months. seasonal patterns, then immediately removing weeds manually as 2. True epiphytes; often chronic; usually filamentous algae; soon as they appear in order to prevent their reproducing and attach to cortical layer; damage plants (Ask, 1999). spreading (e.g. Ask, 1999). 3. Microalgae such as diatoms that form a "scum" on the crop. Removed pest weeds should be taken to land and used as fodder or Often a due to poor water quality (low flow; high silicon; compost material. In the case of epiphytes and epizoa, however, eutrophication) there is little choice but to crop out old stock and replace it with clean propagules.
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5. W
XBC
Good as Gold
Gold The plants shown in the four photos to the left are in condition Gold 1. They have excellent colour density, no grazing and very clean thalli. The plants shown below and to the right are Gold 2 plants. They appear to be in good condition but pigmentation is weakening. These plants must be watched to see whether they are trending toward bleaching (condition yellow).
GOOD AS GOLD - Plants are healthy, with good colour and no more than trace amounts of grazers or W.E.E.D.
Crops that look like the one below are the rule rather than the exception in many places where eucheuma seaplants have been grown for several years - even decades. One of the secrets to this success is that the carrying capacity of local environments not be reduced by poor farming practices.
Environmental impact Some areas appear to enter a period of poor growth after several years of seaplant farming. These usually appear to be places that are very heavily planted - usually using "off bottom" methods that close to sea floors that have been subjected to considerable direct, secondary, collateral or indirect damage. Such places may recover after a few fallow years but should be replanted using less environmentally stressful agronomy practices.
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5. X Monitoring the Environment

Site selection and crop logging activities can only be effectively undertaken if a sound, complementary environmental monitoring program is maintained. The measurements resulting from environmental monitoring can be stored in databases and used in correlation studies among crop-log data and environmental parameters. In many suitable seaplant farming areas a variety of institutions maintain environmental data bases that can be available to seaplant farmers on reasonable terms. Data products of particular use for farm siting, planning and management include the following: 1. Tide tables available form government or international sources or from online programs such as JTides (freeware). 2. Nautical charts, topographic charts, aerial photographs and satellite imagery. 3. Meteorological records from nearby airports, government research stations or weather stations. 4. Oceanographic records from nearby government institutions or private factories and utilities.
Measuring nutrients directly can be costly and tedious so by far the most effective way to track these is to utilise crop-logging parameters such as plant colour indices. The speed and velocity of water currents can be measured using a variety of devices but Doty (1971a) devised a "clod card" system that attaches uniformly-made, appropriately-designed masses of calcium sulphate to wing-like cards. This enables comparison of total water movement among a variety of habitats. Diffusion of calcium sulfate, which is only slightly soluble, provides a measure of water movement that approximates conditions encountered by living seaplants. Salinity checks can be done using a hand-held refractometer (upper right). Temperature records can be maintained on the basis of measurements made using simple minimum-maximum thermometers such as the one shown lower right. Wind speed and direction can be recorded using instruments available from several suppliers (example lower, middle). Sunlight can be monitored using the classic Campbell Stokes heliograph (left) or more modern transducer/data logger combinations such as the Kipp & Zonen unit (below).
Whether or not useful environmental data can be acquired from third parties it is almost always useful for seaplant farmers to undertake local monitoring programs whether individually, as groups or within government, NGO and privately sponsored programs. The most crucial parameters to monitor are water motion, water temperature, wind speed, wind direction, rainfall, solar radiation, salinity and nutrients. Some useful monitoring devices and methods are described opposite.
Right: Manufacturers' photos
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5. Y Monitoring the Crop Situation

Regular crop monitoring programs should include the orderly maintenance of a database that includes plenty of photographs for immediate and future reference. Example forms are shown below: Chart of Scales for Kappaphycus Crop Logging Bleaching Grazing W.E.E.D.
noticable significant severe noticeable or worse low noticeable significant low noticeable significant trace or none noticeable significant severe low noticeable significant low low noticeable significant severe
"A picture is worth a thousand words" - This is a truism never more appropriate than when applied to crop reporting. Aerial photographs give the best overviews but farm over views taken from the ground (e.g. at right) can give an excellent picture of how crops are doing if they are complemented by closer shots such as the photos below.
Code
yellow cyan
# 3 2 1 2 1 3 2 1 2 1 2 1 3 2 1 3
action
scale 1 2 3 4 5 6 7 8 9 10
vigilance move bailout move bailout vigilence move bail out move bailout
noticeable or worse
tan
noticeable or worse slight none little or no bleaching
pink gold
maintain 11 maximise 12 vigilence 13 move bail out move bailout move bailout 14 15 17 18 19 20
low noticeable significant severe
blue
vigilence 16
little or no bleaching
brown
2 1 2 1
It is useful to photograph plants at the extreme range of variation from a site both in close-up (top, left) and at the whole-plant level (below, left). It can also be useful to photograph against a background that has a colour index and ruler for scale (below).
little or no bleaching
purple
Weekly Crop-Log Form

Site name: Week no. Date: Observer: Crop scale at week start: at week end: Km. line: Km.planted: Km.harvested: Kg.planted: Kg.harvested: Photo index ##: Sun Min/max Temp (oC) Rain (mm) Wind direction/ Max speed (m/sec) Mon Tue Wed Thu Fri Sat
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6.A Managing Propagules on the Farm

Once cultivar strains have been selected the key elements of propagule management are: 1. 2. 3. 4 5. Choice of cultivar strain(s) to be grown. Size of propagules to be planted. Age and/or size at which propagules are to be harvested. Spacing of propagules on and within habitat systems. Selection and cutting of propagules for replanting
With respect to the combined effects of propagule size and cropping cycle two distinctly different strategies have been evolved by eucheuma seaplant farmers: 1. Small size - Long cycle with propagules of about 50 - 150 grams cropped at 45-60 day cycles. 2. Large size - Medium cycle with propagules of about 150-300 grams cropped at 30-45 day cycles.
The choice of strain varies with local experience and several are now under cultivation.
Strategy choices are determined by a combination of local conditions and farmers' opinions. In general choppy waters tend to favour "Small-Long" strategies but smooth waters with strong currents The predominant choice for most Kappaphycus farmers seems to be favour "Large-Medium" approaches. In all cases a major determining K. alvarezii var. tambalang of Philippine origin. The Bali strain of E. factor in strategy choice is the point at which significant propagule denticulatum appears to be widespread but several locally developed breakage losses are incurred. Eucheuma strains have been developed in various spinosum Spacing of propagules varies widely among farm regions. producing areas. There is wide variation among farm areas with respect to propagule size at planting and harvesting. Age at harvesting is most commonly set at 40-50 days. In general farmers adopting Small-Long strategies tend to space propagules close together on lines (10-20 cm.) while those adopting Large-Medium strategies space plants more widely on lines (20-30 cm.). Spacing among lines follows a similar trend with the space between lines often being similar to the spacing of plants on lines. This depends partly on space availability, habitat type and currents. Some longline methods involve the spacing of lines several metres apart as a means of reducing line tangling. Selection of clean, vigorous growing tips for replanting is an essential function of farm management; these must be securely attached but not too tightly bound. Generally propagules are broken from their mother plants by transverse cuts or breaks. Trono and Ganzon-Fortes (1989) proposed that slicing plants obliquely yields higher growth rates than transverse cutting. Apical tissue grows faster than basal and median fragments (Mairh et al., 1995). Well branched, obliquely sliced propagules with numerous tips appear to be best for replanting.
Significantly longer and shorter cycles are encountered in some regions. In some areas "pruning" is done as propagules are left attached to lines for many weeks and growing tips are removed. This method may cause quality problems related to an unfavourable mix of young and old tissue in the crop and it is discouraged by many buyers.
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6.B Tender, Loving Care (the TLC Factor)

A superficial examination of eucheuma seaplant farming can mislead the observer into thinking that it is a low-effort occupation such that farmers simply tie cuttings to strings, go away and return to harvest the crop after 5-6 weeks.
The best fertiliser is a farmers shadow on the field a time-tested principle of terrestrial agronomy and seaplant farming. TLC in action (top) and absent (bottom). Same time, same place, same propagule age & source but different farmers (Bali ca. 1990).
Basarun bin Kasim and Arina Ajok TLC experts from Malaysia
Nothing can be further from the truth. Seaplant farming is an occupation such that the most successful are those with skill, diligence and a "green thumb". These attributes must translate to "tender, loving care" (TLC). This means that the farmer must ensure daily attention to functions such as: 1. replacing loose or weak propagules. 2. shaking silt or other loose "scum" off the plants. 3. removing drift material such as plastic bags, debris and weeds that get tangled in the crop. 4. re-attaching or tightening detached or loose lines. 5. replacing or repairing loose netting, stakes, floats, etc. Environmental impact Care for farm structures and crops is essential for success but it is equally important to take care of the surrounding environment. Farmers must take care to avoid trampling or damaging local habitats; littering the environment with trash; polluting farm areas with human and other waste; or undertaking collateral activities such as the use of fish bombing and other destructive activities.
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6.C
Moving Propagules Among Locations
Eucheuma seaplant strains have been dispersed by human activity Once eucheuma seaplants are farmed in the ocean breakage is to the point where marine out-plantings have occurred in at least 33 inevitable and fragments fall to the sea floor. Many are eaten by grazers or fall to areas where they quickly die. countries or territories. Some survive and propagate as loose populations. Broken or cut In some cases dispersion has been within the natural range of Kappaphycus thalli can fuse to each (lower left) or to stones (lower cultivated species but there have been several recorded instances right) so attached populations can occur and in certain conditions may (and no doubt many unrecorded instances) where species have been propagate vegetatively or by sporulation. transplanted beyond their natural ranges. Eucheuma seaplants are robust and easily transported if they are kept moist with seawater and are held within a temperature range that does not harm them (ca. 10-25oC). Under such conditions propagules can live several days and amounts on the order of tens of grams can be propagated to yield thousands of tons of material that form the basis of a regional industry. For the most part transplantations of this sort have been done without benefit of quarantine procedures. Such were the cases of Indonesia (Adnan & Porse, 1989), the Maldives (De Reviers, 1989) and Tanzania (Lirasan & Twide, 1993). In other cases quarantine procedures have been followed as in the Solomon Islands (Smith, 1990), Brazil (De Paula et at., 1998) and India (Mairh, pers. comm.).
Ask et al. (2001) have proposed procedures based on FAO guidelines and specifying that quarantine facilities should have these Zemke-White (in press) has reviewed the literature pertaining to impacts of introduced eucheuma seaplants. He points out that few characteristics: adequately structured studies have been done on the subject and that 1. Isolation from other aquaculture facilities. results so far are inconclusive. It is clear, however, that Hawaii is an 2. Structures that prevent entry of marine organisms. instance where Kappaphycus populations appeared to have little impact at first and have spread to the point of having noticeable 3. Water supply of good quality and independent source. impacts (e.g. covering coral) more than 25 years after their 4 Discharge systems that effectively prevent the inclusion of introduction. biological material. Plants should be maintained in such facilities for at least two weeks Environmental, legal, political and social impacts and should be examined at least daily to check for growth of The pros and cons of transplanting useful species to non-native sites organisms on the thalli. Water should be changed at least twice per for commercial cultivation is a contentious subject. Seaplant farmers week and waste water discharged in a way that ensures no escape must take into account the potential environmental, legal, political and of aquatic organisms to local waters. social consequences of any such introductions. When done introductions should be implemented with proper quarantine precautions.
Kappaphycus alvarezii has been reported by Jennifer Smith and others to be an "alien and invasive" species in Hawaii.
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6. D
Harsh Environmental Conditions
Typhoons and seasonally harsh weather prevent sustained economically viable seaplant farming in some tropical areas or restrict farming to certain seasons. This location in India is calm enough for seaplant farming during the NW monsoon but has wave action such as this during the SW monsoon.
On the day after Typhoon Bising in the Bohol region of the Philippines more than 100 eucheuma seaplant farms and farmhouses had totally disappeared including the ones shown below (author, pers. obs.).
Drifting logs and misdirected boats can damage farms and all manner of flotsam and jetsam can foul floating gear. Farmers cannot anticipate events such as these and there is little that they can do to prevent such random damage. Once destruction by unpredictable events is discerned repairing the broken lines, gathering and reattaching the broken thalli and harvesting those not used in restoring the planting are usually the most economic responses. To some extent gear damage can be minimised of prevented by proper design and construction. For example this type of raft minimises tangling of flotsam and jetsam in the lines.
At least 40 farmers disappeared during this typhoon. See Lim (1982) regarding the damage wrought by this typhoon. In this and other cases the author has noted several instances when farmers were actively repairing and replanting their farms on the day after such devastating events.
Environmental impacts Above all seaplant farmers must treat the forces of nature with respect. Many people lost their lives during typhoon Bising because typhoon warnings were unheeded in the face of previous "false alarms".
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6.E Farm Productivity

Traditionally farm productivity is referred to in terms of production per unit of area. For the vast majority of eucheuma seaplant sea farms, where monolines are used to suspend plants in water, productivity is best regarded as a function of biomass produced per length of line. Length of line per unit of area will then give a unitarea production figure if such is desired. There are several reports of eucheuma seaplant productivity in the literature. Generally daily specific growth rates of 2-6% are cited for commercial farms. Table 6.E below shows the sorts of growth rates that can be expected during good weather conditions with a strain such as K. alvarezii var. tambalang grown using four exemplary techniques:
Km. of line per hectare A B C D 25 25 5 5 Wet 4Dry 4Specific Seedling week week Growth weight yield as yield as Rate (grams) kg/km of kg/km of (%/day) line line 1.5 3 3 4.5 100 200 200 400 260 1,315 1,315 5,060 26 132 132 506 Dry 4week yield as tons per hectare 0.7 3.3 0.7 2.5 Dry tons per hectare per annum 8 43 9 33
These calculated figures are well within the bounds of realised commercial production that the author has observed in several locations. The examples shown represent planting strategies that have been observed in Sabah, Malaysia. Note that some high area production rates (e.g. 33 and 43 dry tons/ha/yr) are postulated but such rates have been met or exceeded several times in the author's experience. The lower rates shown (e.g. 8-9 dry tons/ha/yr) are conservative.
Situation A - Floating or off-bottom system in expansion mode with small seedlings being split and dispersed through the plantation until it is fully planted. Situation B Floating or off-bottom culture in a good location and well tended by the farmer. Situation C Well-tended long line system in a good location with widely spaced lines. Situation D Long line system in an excellent location with fast flowing water.
The compounded growth rates of eucheuma seaplants can lead to rapid farm development and expansion as has been demonstrated at many sites throughout the regions now in commercial production. For example about 200 tons of planting material can be developed from a 20 gm plant growing at an average of only 2.5%/day for 2 years if propagules are constantly replanted (Figure 6.E above).
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6. F Washing and Chemically Treating the Crop

In some areas harvested, live seaplants are washed in seawater to remove silt, salt or other contaminants (photo right).
In Madura, Indonesia farmers speed up drying during rainy spells by utilising a salting" method. Some buyers will not purchase salted weed; others will. It generally brings a price lower that that of unsalted material. This method is similar to salting of fish. It involves the placement of live seaplants into cement vats (below). For every 1,000 kg. of fresh crop about 50 kg of sea salt is added. No water is added. After overnight soaking the crop is removed and sun dried (upper right). The crop bleaches as it dries.
A vermillion coloured saline In other areas they are soaked in weak alkali (about 0.1N) for several "soup" is left in the vat (lower right). hours to "stabilise" the crop . To this is added Alkali stabilisation removes colour and effectively preserves the 1,000 kg. of fresh quality of Eucheuma spp. if properly applied. If the correct crop and 25 kg. combination of soaking and drying conditions is achieved the alkalisalt for a second stabilised "spinosum" can be ground, blended and utilised as a form batch. After that of "processed eucheuma seaweed" (PES) or "semi-refined the soup is carrageenan" (SRC). discarded There is a growing trend for eucheuma seaplants to enter international commerce in the form of "cottonii chips" or "meal". In many cases this is fully alkali-modified material that can be directly ground and blended for sale as PES (E407a) or dissolved in water, clarified and sold as carrageenan (E407). The "chips" process generally involves drying the crop to a moisture content of less than 50%; cooking at 80-90oC for 2-3 hours in the presence of a 1-2N KOH + KCl mix; washing in several volumes of fresh water; chopping; drying; and grinding into coarse "chips" or meal. The end product is generally packed in sacks for shipping to further-processors. Monograph on the way Look for the up-coming SuriaLink Monograph entitled "The ABC of Processed Eucheuma Seaweed - the Semi-refined Carrageenan of Commerce" Environmental impact Processes for washing and treating eucheuma seaplants can yield effluents high in colour, odour, B.O.D., alkali or salts so they must be disposed of with care.
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6. G Drying the Crop

Eucheuma seaplant crops are generally dried before shipment to further-processing facilities. "Industry standard" is about 38% moisture. Individual transactions may involve specifications as low as 30% or as high as the trading environment permits. Attempts at finding better and more cost effective all-season drying options are a persistent industry preoccupation.
Some common means of drying are shown below. These systems usually involve the use of tarpaulins or plastic sheeting that can quickly be used to cover drying crops in the event of rain.
In the photo above the author (left), Tita Barriga Tomayao (centre) and Ruben Barraca (right) puzzle over one of the many systems that has been tried over the years; Cebu, Philippines, 1979.
Wooden platforms over the sea (upper left); concrete slab (above); portable drying flakes made of woven bamboo (left); hanging lines to dry (lower left) tarpaulins or netting over sand or gravel substrate (lower right).
Very well dried Kappaphycus has KCl salt crystals evident on the exterior of plants (lower left) and all well-dried eucheuma seaplants have a stiff texture. The KCl is useful for further processing but some Some farmers dry crops directly on sand or dirt but this practise is buyers prefer to have salt and sand removed (e.g. by flailing, above not popular with buyers because they must then remove sand and right). soil in order to get an acceptable product.
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6. H Cleaning and Packing the Crop

Before they are packed dried eucheuma seaplant crops are usually sorted by hand to remove "junk weeds", raffia and other debris (photo below).
Bales are generally made at weights of about 50 or 100 kg. and are sized to permit loading of 20 tons/20-foot container. Manual screw presses were once used but today bales are usually compressed using hydraulic presses. A wide variety of designs is available.
Following sorting and cleaning the seaplants are tightly packed in sacks (below) or they are compressed into bales (photos opposite)
Sacks of eucheuma seaplants are commonly shipped within Southeast Asia in vessels such as the Filipino "kumpit" (right with 80 tons of seaweed)
Bales may be wrapped and strapped for shipping but some customers prefer "naked bales" with no strapping material at all so they can dump entire container loads and treat them as single raw material lots.
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6. I Quality Testing and Material Balance

Depending on cloud/rain conditions and plant density eucheuma seaplants can typically be dried in 2-3 days under tropical conditions. Plants must be turned over several times per day. The wet : dry ratio of eucheuma seaplant crops varies widely among species and locations. Generally they range from about 6:1 to 9:1. A typical drying curve is shown below: A Moisture test for dried eucheuma seaplants
Materials: 1. Analytical pan balance (accurate to 0.01 gm.) 2. Drying oven 3. Aluminium trays
Procedure: 1. Tare aluminium tray and record weight. 2. Fill tray with 25 gm. sample (should be cut into 3-5 cm. pieces); spread sample thinly and evenly. 3. Dry at 600C (for spinosum) and 800C (for cottonii) for 18 hours. Don't load the oven with materials of widely variable moisture content. 4. Weigh tray and dry weed. Record weight
Calculations: Wt. of dry weed = (wt. of tray + dry weed) - (wt. of tray empty) % MC = (25 gm. - wt of dry weed x 100)/ 25 gm.
A Salt, Sand & salt-free dry matter (SFDM) test

Materials: 1. Analytical Pan Balance (accurate to 0.01 gm.) 2. Drying Oven 3. Aluminum trays 4. Beaker, 4 litre 5. U.S. Standard Sieve No. 12 6. Water Sprinkler 7. Distilled or deionized water Procedure: 1. Weigh accurately 60g of dry weed (precut to about 4 cm.) and place into a 4 litre beaker. 2. Add about 2 litres of water. 3. Soak the weed for 30 minutes with agitation every 5 minutes. 4. Drain off water using sieve No. 12 and spray rinse for about 2 minutes. 5. Tare cafeteria tray and record weight. 6. Dry weed on the cafeteria tray and spread very thinly and evenly (one particle thick). 7. Dry at 600C (for spinosum) and 800C (for cottonii) for 18 hours. Dont load the oven with materials of widely different moisture content. 8. Weigh tray and washed anhydrous weed and record weight.
Basic equipment needed for eucheuma seaplant quality testing (see opposite) includes a good balance and a recirculating drying oven (examples in photos below).
Calculations: % SFDM = (wt. tray + wt. SFDM) - wt. tray empty x 100)/ 60 gm. % Sand & Salt = 100 - (%SFDM + %MC)
There is no generally recognised quick, simple way to measure moisture in eucheuma seaplants. Quick-dry systems tend to incinerate tips and leave thick parts of fronds un-dried. They are useful with powders but not with raw weed. Electronic testers designed for grains (e.g. rice testers) do not function because high seaplant salt levels render readings useless.
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monograph index Page 70

Effective action
6. J
Control Systems
next
As of 2003 an overwhelming percentage of eucheuma seaplant farms Control function Perceptions & information are operated by families in rural areas of tropical countries. In family Real-time functions farming systems crop control is usually effected entirely by human Process control Human, electronic & action based on the collective memory and experience of local mechanical sensors farmers. It must be recognised, however, that farmers are at the base of a supply chain that includes industry and government managers that are also stakeholders with influence on overall supply chain management (Figure 6.J below).
Effects of control action
Monitoring & diagnosis Decision support Strategic & financial Farmer experience, planning databases & simulation models Site selection Geographic, meteorological, oceanographic data Monitoring & Human, electronic & warning mechanical sensors; data & models Legal & regulatory advice Scheduling & planning Logistics & marketing advisory
Manipulation of crop "Smart" operation of systems & control farm systems, devices machinery & equipment Human, electronic & Data, advisories & Guidance for field mechanical sensors warnings operators Reports describing & Management makes rating alternative critical decisions scenarios based on good info Maps & reports Optimum sites indicating degrees of selected for farm suitability placement Actions that reflect Effective & timely warnings & actions that avoid suggested actions perceptible problems Written & codified Warnings & Compliance with copies of laws & suggested actions; complex laws and regulations ready-to-use forms operations Data on weather, Implementation of Anticipation & holidays & crop schedules appropriate reaction "seasonality" to seasonal events Data on market & Optimised Marketing & shipping conditions; marketing & logistic shipping optimised supply prices & plans, routes & to maximise profits conditions schedules Crop logging, Diagnoses, Detection, biological & health prognoses & prevention & cure of data suggested weed, pest & treatments disease problems Cumulative Interactive training; "Hands on" information from simulation trials experience leads to operations & experts useful work
Overall management of this supply-chain involves: 1. Real-time systems that result in immediate effective actions in response to immediately available information. 2. Decision support systems that provide timely information, analyses and advice to system managers and operators. The table opposite summarises control functions typical of seaplant supply chains (after Neish & Ask, 1996). As supply chains evolve purely human controls are being augmented by the tools of technology (Neish & Ask, op. cit.)
Diagnosis of weed, pest & disease events Training
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6.K
Environmental Impacts
Environmental impact notices are presented throughout the present monograph in boxes with this format. Categories of impact relevant to eucheuma seaplant farming are as follows:
Main impacts of farming practices (Zemke-White; in press)
1. Disruption of benthic community structure by removal of macro-benthic organisms and cutting of sea grasses. 2. Substrate abrasion and disruption caused by crops coming into contact with the sea floor. 1. Primary impacts caused by the effects of eucheuma seaplant metabolism and demography. 3. Skewing of species composition caused by the introduction of new sets of ecological niches due to the physical presence of 2. Secondary impacts caused by wastes or other impacts from postseaplants and habitat structures. harvest treatment and handling of crops. 3. Collateral impacts caused by human activities that are directly Zemke-White suggests that farming practices with definite related to seaplant farming including the installation of habitat negative impacts on the local environments include farm refuse systems; trampling of the sea floor; and damage caused by the that litters the environment and tying of anchor lines to live corals. use of boats and vehicles. Other possible negative impacts include shading of underlying 4. Indirect impacts caused by the non-seaplant farming activities habitats; structures built on coral reefs; changes in sedimentation; of farmers and associated species locating domiciles and work and improper treatment of waste water from production facilities. places near to seaplant farms. Possible positive impacts could include increases in fish numbers;
The use of well designed floating systems can minimise impacts of eucheuma seaplant farming because: 1. New habitat is created rather than existing benthic habitat being interfered with. Substrate is placed into the water column where nutrients are most available. 2. Benthic communities are left intact. Planting eucheuma seaplants on or near the sea floor disrupts natural benthic communities and effects species diversity. 3. Crops can be tended using minimally destructive methods. The use of vessels, rafts and dive gear prevents trampling of benthic habitats and organisms.
replacement of destructive activities by farming; and farmers gaining a sense of stewardship over coastal areas. Impacts with either positive or negative effects are changes in primary production; and farms changing the nitrogen regime of the reef community. With respect to species introductions Zemke-White points out that adverse impacts from accidental algal introductions are quite well documented but there have been few studies on the intentional introduction of seaweeds for culture despite the fact that Kappaphycus has been introduced to 19 countries and Eucheuma has been introduced to 13.
While quarantine procedures have been researched, they have been implemented in only two cases before introduction of Kappaphycus Zemke-White (in press) has comprehensively reviewed environmental impacts caused by seaweed farming in the tropics. He new locations. Kappaphycus is well known to escape from farms points out that the farming of eucheuma seaplants is by far the most and can establish free living populations. Impacts of such populations extensive form of tropical seaplant farming. He suggests three main on local habitats may differ between locations but there is evidence from Hawaii that Kappaphycus is overgrowing and killing endemic impacts of farming practices. All are most strongly associated to corals. on/off-bottom methods.
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6.L Eucheuma Seaplants in Multiculture and Polyculture Systems

Theoretically the integration of eucheuma seaplant cultivation into integrated, multiple-plant systems (multiculture) or seaplant/animal systems (polyculture) makes good sense. In practise such systems have been slow to develop beyond the experimental stage. As technology evolves multiculture and polyculture systems may develop further. Seaplant multiculture systems are those in which two or more seaplant species are cultivated by the same enterprise in the same space. The mixed cultivation of Kappaphycus and Eucheuma on the same farm has been commonplace since about 1978 when culture technology for both genera had become firmly established. Multiculture systems involving eucheuma seaplants and agar-bearing seaplants such as Gracilaria have been successfully tried at an experimental level but there has not been widespread development of such systems. Inter-cropping of eucheuma seaplants with other seaplant genera has been tried as a grazer control method with uncertain results (author, pers. obs.). There are also periodic initiatives to make a virtue of necessity by harvesting eucheuma "weeds" such as Ulva and turning them into a companion crop of the eucheuma seaplants. Seaplant/animal polyculture systems were reviewed by Neish and Ask (1996) in the context of expert systems applications to sustainable aquaculture. They presented the following summary of seaweed polyculture systems: LocationType
S.E. Asia ponds Chile tanks Canada Norway Spain
Eucheuma seaplants have been combined in polyculture systems with filter-feeding molluscs, Penaeid crustacea, abalone and finfish. They have served variously as removers of nitrogenous waste, as substrate, as feed and as companion crops. All systems known to the author have been experimental (e.g. the Kahuku, Hawaii system). The development of multiculture and polyculture systems appears to have potential for yielding the following benefits to seaplant farmers and other mariculturists: 1. Sharing of coastal areas suitable for mariculture. Such areas will become ever more scarce and valuable as mariculture competes with other foreshore activities so sharing is becoming necessary. 2. Sharing of other expensive or scarce resources such as management talent, transportation infrastructure and ocean structures. This can provide economies of scale. 3. Utilisation of synergies and complementary characteristics of crop species to minimize pollution and reduce production costs. 4 Enhancement of economic stability through diversification of the economic base of mariculture operations. Perhaps the lack of development in multiculture and polyculture involving eucheuma seaplants is due in large part to the fact that they are overwhelmingly cultivated on small family farms in tropical areas where farmers gain as much income as they aspire to from their main crop alone (author, pers. obs.). "High tech" eucheuma seaplant systems and multiculture or polyculture systems may become economically significant in the future as the need for more intensive utilisation of marine foreshores develops and as the aspirations of people from sea farming communities rise. When this occurs the modern tools of communication and artificial intelligence will probably play a significant role (Neish and Ask, 1996).
Animals
prawns salmon
Seaweeds
Gracilaria spp. Gracilaria spp.
Reference
Hurtado-Ponce, 1995 Buschmann et al, 1995 Petrell, 1995 Bodvin, 1995 Jimenez del Rio, 1995
sea cages rafts salmon kelp sea enclosures salmon, mussels kelp tanks golden bream Ulva rigida
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Acknowledgements
Maxwell Stanford Doty
Arthur Charles Neish
This monograph is dedicated to the author's two major seaplant mentors; Dr. Arthur Charles Neish and Dr. Maxwell Stanford Doty. Neish and Doty were contemporaries and both did pioneering work on the cultivation of red algal galactan seaplants. During the late 1960s and early 1970s Neish was busy working on Chondrus in the Canadian Atlantic while Doty was engaged in developing the cultivation of eucheuma seaweeds in the Indo-Pacific region. The efforts of both of these seaplant pioneers contributed to the commercial cultivation of seaplants; to the acquisition of a great deal of seaplant knowledge and to the development of a good many seaplant people for whom Max and Art became "gurus".
My debt to Arthur Neish is considerable not only because he was my father but also because we became scientific colleagues during the latter years of a life that was prematurely ended by occupation-related cancer. Me, my brothers Gordon and Doug and my sister Nancy all learned the foundations of science from our parents at the dinner table or during the excursions to "the lab" that was sometimes our father's idea of entertaining the kids. In its small way, the SuriaLink site, this monograph and my seaplant career are all attempts to continue some of the work that my father started. "Information Technology" was in its infancy when my father died but he would have enjoyed its bounties immensely. The SuriaLink site has developed from the system of research bulletins that my father developed as a means for getting information disseminated quickly while he was with NRCC (e.g. Neish & Shacklock, 1971).
Besides his considerable debt to Doty and Neish the author is very much indebted to many people who have taught him about seaplant farming and biology over the past 35 years. These include the late Vicente Alvarez, Erick Ask, Ruben Barraca, Made Simbik, Basarun bin Kasim, Farley Baricuatro and several other former colleagues from Marine Colloids (later FMC); also numerous seaweed farmers and Last - and certainly not least - I give thanks to my good friend seaplant business people who I have met over the years from all over and colleague Dr. David Myslabodsky for exchanging ideas, for the world and people who contact me through SuriaLink. reviewing the manuscript and for contributing useful material to the sections of this monograph that involve chemistry.
My debt to Max Doty originated as a result of correspondence between him and my father. Max's counsel was of inestimable value to me as I ascended the learning curve for the eucheumas and other tropical seaweeds. I used to stop off in Hawaii frequently during the period 1977-1997 and until he passed away Max was at the top of my list of people to meet during my time there. The present monograph is very much an outgrowth of the discussions we had. If Max had lived longer we had hoped to co-author a monograph covering this subject matter and no doubt it would have been far richer from his contribution.
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Glossary
Agar Agronomy Apical Axenic Basal Biopolymer Callus Carpospore Carrageenan Cisternae Clone Conjugate Cortex Cultivar Dioecious Diploid Frond Furcellaran Gamete Gametophyte Germinate Golgi body
Red algal galactan biopolymer produced by genera such as Gracilaria, Gelidium and Gelidiella. The arts and sciences of managing crops on farms Pertaining to the terminal segment or "tips" of fronds. Uncontaminated and germ free (applied to cultures) Pertaining to the oldest segment or "base" of fronds. Compound of high molecular weight synthesised by living organisms Tissue that forms over cut parts of fronds Diploid spores produced by carposporophytes that live parasitically on their mother plants Red algal galactan biopolymers produced by genera such as Kappaphycus, Eucheuma, Betaphycus, Gigartina, Chondrus and others. Reservoirs or receptacles that hold fluid in the plant tissue A group of organisms derived from a single individual Fusion of two one celled organisms for reproduction where fertilisation occurs The pigmented outer cell layer of a thallus or frond A genetically uniform set of propagules Organisms that have male and female reproductive structures on different individual members of the species Having two similar complements of chromosomes A branch of a thallus Red algal galactan biopolymer produced by Furcellaria spp. Mature haploid reproductive cell capable of fusion with another gamete to form a diploid nucleus Life cycle stage in many plants and algae; individual plant composed of haploid cells that produce gametes To begin growing or developing Golgi apparatus/body; a net-like mass of material in the plant cytoplasm that is a site of biopolymer synthesis
Habituate Haploid Macroalga Medulla Microalga
To become used-to or adapted-to stimuli Having one complement of chromosomes Non-vascular aquatic or marine plants of the phyla Chlorophyta, Rhodophyta and Phaeophyta; Large enough to be seen using the naked eye The un-pigmented cell layer immediately below the cortex Non vascular aquatic or marine plants too small to be seen by using the naked eye. on the same individual
Macrophyte Plants large enough to be readily seen by the naked eye
Monoecious Organisms that have both male and female reproductive structures Morphology Form and structure of the plant Pericarp Phenotype
The walls of a ripened fruiting body A character or individual defined by its appearance and not by its genetic makeup alginates and carrageenan)
Phycocolloid Complex polysaccharide biopolymers produced by algae (e.g. agar, Phycologist A scientist who studies algae Propagule Protoplast Rhizoid Seaplant Seaweed SFDM SGR Sporophyll
A cutting or fragment of a seaplant thallus that is used for vegetative propagation of a crop Actively metabolising membrane-bound part of a cell (as distinct from the cell wall). Root-like filaments by which a macroalga attaches to substrate; collectively may form a holdfast Any photosynthesising organism that lives in seawater Common name applied to most marine macroalgae Salt free dry matter Specific growth rate expressed in percent per day Structure that produces reproductive cells called spores produce spores
Sporophyte The life cycle stage in plants and algae that terminates in meiosis to Tetraspore Thallus Uniseriate
one of four asexual spores produced in a tetrasporangium The entire physical entity of a propagule or a whole plant Occurring in a single series
Gonimoblast filaments that extend from the egg cell to support carposporophytes
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Walker, D. I. and Kendrick, G. A., (1998). Threats to macroalgal diversity: Marine habitat destruction and fragmentation, pollution and introduced species. Botanica Marina, 41 (1) : 105-112. Weber-van Bosse, A., 1913. Marine Algae, Rhodophyceae, of the "Sealark" Expedition, collected by Mr. J. Stanley Gardiner. Trans. Linn. Soc., Lond., Ser. 2.,Bot. 8, pp. 105-142. Weber-van Bosse, A., 1926. Algues de l'expedition danoise aux iles kei. Vidensk. Mddr. dan. Naturhist. Foren. 81, pp. 57-155. Weber-van Bosse, A., 1928. Liste des algues du Siboga. IV. Part 3: Gigartinales et Rhodymeniales. In M. Weber (ed.), Siboga Expedie, Monog. 59d, pp. 393-533. Wu, C.Y, Li, J.J., Xia, E.Z., Peng, Z.S., Tan, S.Z., Li, J., Wen, Z.C., Huang, X.H., Cai, Z.L., Chen, G.J., 1988. Transplant and artificial cultivation of Eucheuma striatum in China. Oceanol. Limnol. Sin. 19, pp. 410-417. Yokuchi, A., 1983. Seasonal variation of spore generation and development of tetraspores of Eucheurna gelatinae J.Agardh (Gigartinales, Rhodophyta) at Eriomote Is. Jpn. J. Phycol. 31, pp. 34-37. Yamada, Y., 1936. the species of Eucheuma from Ryukyu and Formosa. Sci. Pap. Inst. Alg. Res., Fac. Sci., Hokkaido Univ. 1, pp. 119-134. Zablackis, E., V. Vreeland, B. Doboszewski & W.M. Laetsch, 1988. Localization of kappa carrageenan in cell walls of Eucheuma alvarezii var. tambalang with in situ hybridization probes. In T. Stadler, et al. (eds), Algal Biotechnology. Elsevier Applied Science, New York, pp. 441-449. Zemke-White, W.L., in press. Assessment of the current knowledge on the environmental impacts of seaweed farming in the tropics. Proceedings of the AsiaPacific Conference on Marine Science and Technology, 12-16 May 2002, Kuala Lumpor, Malaysia. Zemke-White, W. L. & M. Ohno, 1999. World seaweed utilisation: an end-ofcentury summary. Journal of Applied Phycology 11, pp. 369-376. Zertuche-Gonzalez, IA., 1988. In situ life history, growth and carrageenan characteristics of Eucheurna uncinaturn (Setchell & Gardner) Dawson from the Gulf of California. PhD dissertation, SUNYat Stony Brook, Stony Brook, New York, 162 pp. Zertuche-Gonzales, J. A, Z. Garcia Ezquivel & B.H. Brinkhuis, 1987. Tank culture of the red seaweed Eucheuma uncinatum from the Gulf of California. Cienc. Mar. 13(2): 1-18.

The ABC of Eucheuma Seaplant Production

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Monograph # 1-0703, SuriaLink, July, 2003

The ABC of Eucheuma Seaplant Production

1. Abstract & Index

Eucheuma seaplant farm, Bohol, Philippines

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Introduction to the Biology of Eucheuma Seaplants

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Introduction to the Agronomy of Eucheuma Seaweeds

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3.A Taxonomic Classification

Genus: Betaphycus Genus: Eucheuma

Trade name: gelatinae

Authority: Doty ex P.C. Silva Symbol: EU

Type species: Betaphycus philippinensis Doty Authority: J. Agardh

Commercial species: gelatinae (GEL)

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3.B Taxonomic Descriptions

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3.C Table of Eucheumatoideae Characteristics

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3.D Trade Names and Varieties

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3.E Kappaphycus alvarezii var. tambalang (cottonii of the trade)

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M.S. Doty drawing - Length of bar ca. 10 cm. relative to image

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3. G Eucheuma denticulatum (spinosum of the trade)

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3.H Betaphycus gelatinae (gelatinae of the trade)

Gelatinae drying in Hainan, China

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3.I Natural Distribution and Dispersion by Humans

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3.J Commercial Activity

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3.K Commercial Distribution

Country China India Indonesia Madagascar Malaysia Philippines Tanzania Totals

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3.L Commercial Significance

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3.M Commercial Production Trends 1961-2002

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3.O Life History - Overview & Tetrasporangia

M.S. Doty image

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3.P Life History - Male and Female

M.S. Doty image

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3.Q Life History - Carpospores

M.S. Doty image

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3.R Selection of Cultivar Strains

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3.S Varietal Improvement

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3.T Carrageenan Synthesis

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3.U Carrageenan Differences Among Species

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Basic Elements of Seaplant Growth

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