Eucheuma Seaplant Production

SuriaLink Infomedia
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
1. Abstract & Index
Photos and diagrams by Iain C. Neish except where noted otherwise
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
5.Methods
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
6.Crop
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
7.Glossary
8.Acknowledgements
9.Bibliography
Abb-Bod|Bor-Dix|Dot-Gle|Gom-Liu|Lux-Ola|Pad-Sid|Smi-Vre|Wal-Zer
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.
Eucheuma seaplant farm, Bohol, Philippines
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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.
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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.
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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
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This monograph deals with commercially useful genera of the Tribe

Eucheumatoideae; . These are colloquially known as the "eucheuma
seaweeds". Their taxonomic classification is :
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.
Phylum Rhodophyta, Class Rhodophyceae, Subclass Florideophycidae, Order

Gigartinales, Family Areschougiaceae, Tribe Eucheumatoideae
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 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.
Genus: Betaphycus
Trade name: gelatinae
Commercial species: gelatinae (GEL)

Genus: Eucheuma
Symbol: BE
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).
Authority: Doty ex P.C. Silva
Type species: Betaphycus philippinensis Doty
Common names: Eucheuma, Gelatinae (also see Eucheuma names)

Trade name: spinosum
Symbol: EU
Authority: J. Agardh
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
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Eucheumatoideae species of commercial significance are notoriously

variable in form.
Species in section Gelatinae tend to grow in very turbulent, active

water and seem to be unusually tough.
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.
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.
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Characteristics of the typical species in the major sections of

Eucheuma within the Tribe Eucheumatoideae: Gelatinformia and
Anaxiferae.
Eucheuma section
Cottoniformia
Eucheuma section
Eucheuma
Eucheuma section
Gelatiformia
Eucheuma section
Anaxiferae
1- Fronds of many forms but

commonly cylindrical; simple
blunt or spiny protuberances
1- Fronds cylindrical; spines

simple; basal diameters less
than axis thicknesses.
1- Fronds compressed; spines

simple; basal diameters equal
axis margin thickness
1- Fronds cylindrical or
dorsoventral and bearing
compound spines
2- Protuberances irregularly
arranged; in some cases
appearing in vertical rows so
segments become angular
2- Spines in regularly spaced

pairs or whorls first, but later
others may appear scattered
2- Spines in rows, marginally

first and latter dorsally and
ventrally on flatter faces or
scattered
2- Spines often scattered, in

whorls or covering the thallus in
various arrangements or
densities
3- Branching irregular;
sometimes irregularly pinnate;
may be opposite or falsely
dichotomous
3- Branches generally from

whorls; often opposite; irregular
or in pectinate series
3- Branches mostly marginal,

pinnate, often opposite or
irregular but not in pectinate
series
3- Branching generally from

whorls; often opposite; whorled
or irregular
4- Hyphal axial core usually

present; not rhizoidal;
cylindrical.
4- Axial core rhizoidal and

cylindrical
4- Axial core tortuous, hyphal

and often flattened.
4- No hyphal or rhizoidalaxial
core in cylindrical axes
5- Kappa carrageenan
5- Iota carrageenan
5- Beta or other carrageenans
5- Iota carrageenan
6- Cystocarps on main axes
6- Cystocarps on lateral axes
6- Cystocarps on lateral or
pedical axes
6- Cystocarps on main axes
7- No cystocarp associated with

laterals
7- Generally a terminal spine

beyond cystocarp
7- Often several spines on

cystocarp; sometimes none
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.
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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)
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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.
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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).
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.
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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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)
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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
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Natural stocks of Eucheuma seaplant species occur naturally

throughout the Indo-Pacific region from eastern Africa to Guam.
Human actions have had a major impact on the distribution and

abundance of eucheuma seaplants.
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.
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).
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.
next
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
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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.
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.
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.
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.
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.
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).

SuriaLink 1-0703
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3.K Commercial Distribution
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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)
Country
China
India
Indonesia
Madagascar
Malaysia
Philippines
Tanzania
Totals
'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)
Antigua, Barbados, Jamaica, St. Lucia

Belize
Brazil
Cambodia
Cuba
Djibouti
Fiji
French Antilles
French Polynesia
Guam
Honduras
Japan
Kenya
Kiribati
Maldives
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)
ca.1980s
ca.1980s
1995
2000
1991
1973
ca.1978
1978
ca.1985
ca.1985
1978
1983
1996
1977
1986
not known
not known
pre-comm.
pre-comm.
pre-comm.
ceased
pre-comm.
ceased
ceased
ceased
ceased
ceased
pre-comm.
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
trace
trace
nil
nil
nil
nil
nil
nil
nil
nil
nil
nil
trace
nil
nil
static
ceased
static
expand
static
ceased
static
ceased
ceased
ceased
ceased
ceased
static
static
static
+
+
?
+
?
?
+
+
+
+
+
+
+
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)
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.1985
ca. 1990s
2000
ca.1978
1987
1983
1977
1971
1996
1993
pre-comm.
pre-comm.
pre-comm.
pre-comm.
ceased
pre-comm.
pre-comm.
pre-comm.
ceased
pre-comm.
pre-comm.
nil
small/variable
trace
small/variable
ceased
small/variable
small/variable
small/variable
nil
small/variable
small/variable
trace
nil
trace
nil
nil
nil
nil
nil
nil
nil
nil
unknown
static
unknown
expand
ceased
expand
static
static
ceased
static
expand
+
+
+
?
+
+
+
?
+
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)
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)
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)
SuriaLink 1-0703
3.L Commercial Significance

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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)
SuriaLink 1-0703

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).
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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.
SuriaLink 1-0703

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)
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.
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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.
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).
SuriaLink 1-0703

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).
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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.
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.
M.S. Doty image
SuriaLink 1-0703

3.P Life History - Male and Female
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Male structures are seldom seen among eucheuma seaplants except

as reported in Eucheuma isiforme.
In the Eucheumatoideae fertile spine development is important

systematically (Doty, 1988).
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.
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.
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.
M.S. Doty image
SuriaLink 1-0703

3.Q Life History - Carpospores
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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
SuriaLink 1-0703

monograph index
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.
Page 21
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.
next
Above: Colour variants in the harvest at Gunung Payong, Bali,

Indonesia
Left: Tending the crop in Kudat, Sabah, Malaysia.
SuriaLink 1-0703

3.S Varietal Improvement
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To date genetic work by classic hybridisation techniques has not led

to varieties of eucheuma seaplants superior in growth, farmability
or carrageenan characteristics.
Recently Reddy et al (2003) in India have utilised in vitro somatic

embryogenesis and the regeneration of somatic embryos to yield
whole plants.
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.
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.
SuriaLink 1-0703

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.
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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).
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).
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
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Carrageenans are characteristically sulphated with the exception of

"beta" which lacks sulphation and therefore strains the definition of
"carrageenan" (Santos, 1989).
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.
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.
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.
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.
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
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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

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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
Max/range in deg. C
Author or source
Kappaphycus alvarezii
22.8 - 29.2
Ohno & Orosco (1987).
Eucheuma amakusaensis
18 - 25
Ohmi & Shinmura (1976).
Eucheuma denticulatum
Max. 34
Dawes (1979).
Eucheuma isiforme
Max.18-25
Mathieson & Dawes (1974)
Kappaphycus striatum
20 to 30; 22.8-29.2
Mairh et al. (1986); Ohmi

& Orosco (1987).
Eucheuma uncinatum
Max. 24
Dawes (1979).
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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
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Exposure to optimal amounts and wavelengths of photosynthetically

active radiation (PAR) is undoubtedly as essential for eucheuma
seaplants as it is for other plants.
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.
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.
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.
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.
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).
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.
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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.
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.
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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).
Salinity and terrestrial influences are significant for seaplants in both
negative and positive ways.

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4.G Macronutrients
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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
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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.
Water motion NH3-N NO3+NO2-N PO4-P Oxygen
Upstream
45
1.98
1.44
0.66
7.22
Downstream
38
2.01
1.10
0.62
8.25
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.
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)
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).
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.
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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.
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.
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).
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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.
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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.
SuriaLink 1-0703

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.
Mask and fins
rugged GPS
Marine binoculars
rugged computer
water proof Pelican case
manufacturers' photos
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next
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.

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5. A
Agronomy Overview
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.
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Elements Requiring Management

People
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
Development, construction and operation of the physical

structures that comprise the seaplant farm.
Crop
The population of eucheuma seaplants that comprise the farm

populations and yield the crop.
Monitoring functions
People
Semporna, Sabah, Malaysia, 1996
Elements and functions of this process are shown in Fig. 5.A below
and are further described in Table 5.A opposite.
Maintaining useful interaction with stakeholders.
Environment Measurement, recording and data handling for meteorological

and oceanographic parameters.
Habitat
Oversight and inspection leading to preventative maintenance

of farm structures and equipment.
Crop
Crop logging.
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
Operation of farm structures and equipment through manually

applied operational protocols and automated control systems.
Crop
Planting, maintenance and harvesting and post-harvest

treatment of the crop.
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5. B

Some Background
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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.
way to tying cuttings on nets.
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:
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
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.
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

monograph index
Types of Habitat System
The most obvious physical manifestation of seaplant farms is their

habitat structures.
Page 38
Seaplant farm habitat characteristics; list and descriptions:
Substrate or enclosure type
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
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
Position or location
Horizontal net culture, Kappaphycus alvarezii, Sulu Sea, 1977
on
Propagules placed on sea floor
off bottom
Propagules placed just above sea floor
mid water
Propagules placed in water column
surface
Propagules floating at or near sea surface
on land
Farm systems placed in ponds on land; intertidal or above tides
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.
the position or location of the substrate or enclosure relative

to the sea floor and the sea surface.
floats+anchors Support substrate or enclosures in water column or at/near sea surface
3.
whether the substrate or enclosure is oriented horizontal or

perpendicular to the sea surface (or has no fixed orientation).
rafts
Substrate or enclosures suspended from bamboo or plastic-tube rafts
terrestrial
Farm systems placed in ponds on land; intertidal or above tides
4.
the method by which the substrate or enclosure is fixed,

suspended or held in place within the farm area.
Attachment method
none
Propagules unattached; mixed by physical agitation or free floating
the method by which propagules are attached to or

suspended within the substrate or enclosure.
enclosure
Propagules individually unattached but contained in enclosure
tie
Propagules individually tied to substrate using knots
loop
Propagules individually attached to substrate using loops
5.
next

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5. D
Broadcasting and Tie-on-stones Systems
Substrate
Position
Orientation
Fixation
Attachment
coral/stones
on bottom
horizontal
stakes/rocks
none or tie
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
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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
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.
Photos above and below by Jack Fisher, 1977

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5. E
Nets and Monolines in "Off-bottom" Systems
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"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.
Substrate
Position
Orientation
Fixation
Attachment
K. alvarezii
growing on
netting on a
test plot
operated by
Vic Alvarez in
Sabah
(1977). The
"tambalang"
variety
probably
originated
near here.
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.
above: Tanzania, 1991

SuriaLink 1-0703
5. F
Pen, Cage, Sack Bag and Tube Systems
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.
Enclosure
Position
Orientation
Fixation
Attachment
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
Float + Anchor Long-line and Short-line Systems
Substrate
Position
Orientation
Fixation
Attachment
monoline
mid water
horizontal
floats + anchors
tie or loop
net/film bag or tube
surface
perpendicular
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The "long-lines" illustrated below work well where large areas of

water deeper than about two metres are available for farms.
enclosure
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
Raft Systems
Page 43
Substrate
Position
Orientation
Fixation
Attachment
monoline
surface
horizontal
floats + anchors
tie or loop
net/film bag or tube
monograph index
next
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.
enclosure
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
Terrestrial Pond and Raceway Systems
Enclosure
Position
Orientation
Fixation
Attachment
pond/raceway
on land
mixed/agitated
terrestrial
none
net/film bag/tube
enclosure
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.
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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
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).
Experimental polyculture systems that

incorporated eucheuma seaplants (Kahuku,
Hawaii ca. 1981. Jack Fisher shown working
above.)
Agitation involves substantial energy consumption. That tends to

restrict these systems to high-value crops.

SuriaLink 1-0703
monograph index
5. J Positions and Orientations of Habitat Systems

Substrate/enclosure Position
Orientation
Fixation
Attachment
coral/stones
on bottom
horizontal
stakes/rocks
none
pen/cage/sack
off bottom
perpendicular
floats + anchors enclosure
monoline
mid water
mixed/agitated
rafts
tie
net bag/tube
surface
terrestrial
loop
film bag/tube
terrestrial
next
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.
pond/raceway
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.
Page 45
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
Page 46
Substrate
Position
Orientation
Fixation
Attachment
monoline
on/off bottom
horizontal
stakes/rocks
tie or loop
mid water to surface
perpendicular
rafts, floats + anchors
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.
next
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.
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).
monograph index

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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
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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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.
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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.
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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:
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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:
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.
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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5. O Crop Logging Parameters and Actions

AYZ
Pink
Crop logging need not be complicated or technical. Main parameters to monitor are shown in
the paradigm below.
GRAZING + BLEACHING
+ W.E.E.D. - Early
symptoms of bleaching;
noticeable grazing +
W.E.E.D.
AYC
next
Purple
GRAZING + W.E.E.D. Plants healthy; good

colour many grazers +
W.E.E.D
Blue
GRAZING - Plants
healthy; good colour;
noticeable grazing.
Significant grazing
means significant
losses; severe grazing
means grazers
harvesting most of crop.
Cyan
XYC
Tan
W.E.E.D. + BLEACHING Early symptoms of

bleaching; Weeds,
Epiphytes, Epizoa or
Disease.
ABC
XYZ
XBZ
GRAZING + BLEACHING Plants show early

symptoms of bleaching
and show signs of
noticeable grazing.
ABZ
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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.
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.
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
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.
LIME =
MAINTAIN
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.
AMBER =
BE VIGILANT
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

ABC
Yellow
ABZ
Tan
AYC
Cyan
AYZ
Pink
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.
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5. R Types of Grazer Damage

XBZ
Blue
XYZ
Purple
AYZ
Pink
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AYC
Cyan
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.
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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.
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.
next

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5. S Examples of Grazers
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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
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"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
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
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).
XBZ
Blue
XYZ
Purple
AYZ
Pink
AYC
Cyan

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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".
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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).
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.
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
Brown
ABZ
Tan
XYZ
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Purple
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).
next
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
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Gold
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.
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).
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
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Site selection and crop logging activities can only be effectively

undertaken if a sound, complementary environmental monitoring
program is maintained.
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 measurements resulting from environmental monitoring can be

stored in databases and used in correlation studies among crop-log
data and environmental parameters.
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.
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.
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
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).

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5. Y Monitoring the Crop Situation
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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.
Code
action
scale
noticable
significant
severe
low
low
yellow
3
2
1
vigilance
move
bailout
1
2
3
noticeable or worse
noticeable
significant
low
cyan
2
1
move
bailout
4
5
vigilence
low
noticeable
significant
severe
move
bail out
move
bailout
9
10
noticeable or worse
tan
noticeable or worse
noticeable
significant
pink
2
1
slight
none
trace or none
gold
2
1
maintain 11
maximise 12
vigilence 13
little or no bleaching
noticeable
significant
severe
low
low
noticeable
significant
severe
blue
noticeable
significant
move
14
bail out
15
3
brown
purple
vigilence 16
move
17
bailout
18
2
1
move
bailout
19
20
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 ##:
Min/max Temp (oC)

Rain (mm)
Wind direction/ Max speed (m/sec)
Mon
Tue
Wed
Thu
Fri
next
"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.
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).
Weekly Crop-Log Form
Sun
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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
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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.
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.
The choice of strain varies with local experience and several are now
under cultivation.
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.
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.
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.

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6.B Tender, Loving Care (the TLC Factor)
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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).
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.
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.
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
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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.).
Kappaphycus alvarezii has been reported by Jennifer Smith and others

to be an "alien and invasive" species in Hawaii.
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.
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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.
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.
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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.).
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
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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
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Wet 4Dry 4Specific

Seedling
week
week
Growth
weight
yield as
yield as
Rate
(grams) kg/km of kg/km of
(%/day)
line
line
Dry 4week
yield as
tons per
hectare
Dry tons
per
hectare
per
annum
25
1.5
100
260
26
0.7
25
200
1,315
132
3.3
43
200
1,315
132
0.7
4.5
400
5,060
506
2.5
33
next
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).
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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"
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.
SuriaLink 1-0703

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.
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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.
Attempts at finding better and more cost effective all-season drying

options are a persistent industry preoccupation.
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).
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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)
monograph index
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:
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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
Basic equipment needed for eucheuma seaplant quality testing (see

opposite) includes a good balance and a recirculating drying oven
(examples in photos below).
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.
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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6. J

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Control Systems
Page 70
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).
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
Scheduling &
planning
Logistics &
marketing advisory
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.)
Effects of control
action
Manipulation of crop "Smart" operation of

systems & control
farm systems,
devices
machinery &
equipment
Human, electronic & Data, advisories &
Guidance for field
mechanical sensors warnings
operators
Legal & regulatory

advice
Overall management of this supply-chain involves:
Effective action
next
Diagnosis of weed,
pest & disease
events
Training
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
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6.K

Environmental Impacts
Environmental impact notices
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Main impacts of farming practices (Zemke-White; in press)
are presented throughout the present monograph in boxes with this

format. Categories of impact relevant to eucheuma seaplant farming
are as follows:
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
Page 72
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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
Animals
Seaweeds
Reference
S.E. Asia ponds

Chile
tanks
prawns
salmon
Gracilaria spp.
Gracilaria spp.
Hurtado-Ponce, 1995
Buschmann et al,
1995
Petrell, 1995
Bodvin, 1995
Jimenez del Rio,
1995
Canada
Norway
Spain
sea cages rafts salmon

kelp
sea enclosures salmon, mussels kelp
tanks
golden bream
Ulva rigida
next
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).
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Acknowledgements
Maxwell Stanford Doty
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Page 73
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".
next
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).
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.
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.
SuriaLink 1-0703

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Glossary
Page 74 next
Agar
Red algal galactan biopolymer produced by genera such as

Gracilaria, Gelidium and Gelidiella.
Habituate
To become used-to or adapted-to stimuli
Agronomy
Haploid
Having one complement of chromosomes
The arts and sciences of managing crops on farms
Apical
Pertaining to the terminal segment or "tips" of fronds.
Macrophyte Plants large enough to be readily seen by the naked eye
Axenic
Uncontaminated and germ free (applied to cultures)
Basal
Pertaining to the oldest segment or "base" of fronds.
Biopolymer
Compound of high molecular weight synthesised by living

organisms
Callus
Tissue that forms over cut parts of fronds
Carpospore
Diploid spores produced by carposporophytes that live parasitically on their

mother plants
Carrageenan
Red algal galactan biopolymers produced by genera such as

Kappaphycus, Eucheuma, Betaphycus, Gigartina, Chondrus and
others.
Cisternae
Reservoirs or receptacles that hold fluid in the plant tissue
Clone
A group of organisms derived from a single individual
Phycocolloid Complex polysaccharide biopolymers produced by algae (e.g. agar,
Conjugate
Fusion of two one celled organisms for reproduction where

fertilisation occurs
Phycologist A scientist who studies algae
Cortex
The pigmented outer cell layer of a thallus or frond
Cultivar
A genetically uniform set of propagules
Dioecious
Organisms that have male and female reproductive structures on

different individual members of the species
Diploid
Having two similar complements of chromosomes
Frond
Macroalga
Non-vascular aquatic or marine plants of the phyla Chlorophyta,

Rhodophyta and Phaeophyta; Large enough to be seen using the
naked eye
Medulla
The un-pigmented cell layer immediately below the cortex
Microalga
Non vascular aquatic or marine plants too small to be seen by using

the naked eye.
Monoecious Organisms that have both male and female reproductive structures
on the same individual
Morphology Form and structure of the plant

Pericarp
The walls of a ripened fruiting body
Phenotype
A character or individual defined by its appearance and not by its

genetic makeup
alginates and carrageenan)
Propagule
A cutting or fragment of a seaplant thallus that is used for

vegetative propagation of a crop
Protoplast
Actively metabolising membrane-bound part of a cell (as distinct

from the cell wall).
Rhizoid
Root-like filaments by which a macroalga attaches to substrate;

collectively may form a holdfast
A branch of a thallus
Seaplant
Any photosynthesising organism that lives in seawater
Furcellaran
Red algal galactan biopolymer produced by Furcellaria spp.
Seaweed
Common name applied to most marine macroalgae
Gamete
Mature haploid reproductive cell capable of fusion with another

gamete to form a diploid nucleus
SFDM
Salt free dry matter
Gametophyte
SGR
Specific growth rate expressed in percent per day
Life cycle stage in many plants and algae; individual plant

composed of haploid cells that produce gametes
Sporophyll
Structure that produces reproductive cells called spores
Germinate
To begin growing or developing
Sporophyte The life cycle stage in plants and algae that terminates in meiosis to
Golgi body
Golgi apparatus/body; a net-like mass of material in the plant

cytoplasm that is a site of biopolymer synthesis
Tetraspore
one of four asexual spores produced in a tetrasporangium
Thallus
The entire physical entity of a propagule or a whole plant
Uniseriate
Occurring in a single series
Gonimoblast filaments that extend from the egg cell to support carposporophytes
produce spores
SuriaLink 1-0703

Bibliography Abb-Bod
Abbott, I.A. (ed.), 1988. Taxonomy of Economic Seaweeds II. California Sea
Grant Publication.
Abbott, I.A. & J. N. Norris (eds), 1985. Taxonomy of Economic Seaweeds.
California Sea Grant College Program, Univ. of California, La Jolla, California,
U.S.A., 167 pp.
Adnan, H., Porse, H., 1987. Culture of Eucheuma cottonii and Eucheuma
spinosum in Indonesia. Hydrobiologia 151-152, 355-358.
Agardh, J.G., 1847. Nya alger fran Mexico. Ofvers. K. Vet. Akad. Forhandl. 4:517.
Agardh, J.G., 1852. Eucheuma. Pp. 624-629 in: Species, genera et ordines
algarum, Vol. 2(2). C.W.K. gleerup, Lund, pp. 337-720.
Agardh, J.G., 1876. CLI: Eucheuma. Pp. 598-603 in Species, genera et ordines
algarum 3(1): Epicrisis systematis Floridearum. Leipzig, 724 pp.
Agardh, J.G., 1892. Observationes de speciebus algarum minus cognitis
earumque dispositione. Pp. 118-126, in Analecta Algologica. Lunds Univ. Arsskr.
26(3): 1-182.
Aguilana, J. T., Broomb, J.E., Hemmingson, J.A., Dayrita, F.M., Montao,
M.N.E., Dancel, M.C.A., M. R. Nionuevoa, M.R. & Furneaux, R.H. 2003.
Structural Analysis of Carrageenan from Farmed Varieties of Philippine Seaweed,
Botanica Marina Vol. 46, 2003, pp. 179192.
Alih, E.M., 1990. Economics of Eucheuma farming in Tawi-Tawi island in the
Philippines. In: Hinano, R., Haayu, I. (Eds.), Proceedings of the 2nd Asian Fisheries
Forum, Tokyo, Japan, 17-22 April, 1989, vol.2, pp.249-252.
Amat, J. P., 1981. Etude et Culture de l'algue Rouge Eucheuma spinosum dans le
Golfe de Tadjourah. Thesis, l'Institut National de Toulouse, 125 pp.
Anderson, G. W., 1953. A note on the seaweed resources of Zanzibar
Protectorate. Proc. int. Seaweed Symp. 1: 102f.
Ask, E.I., 1999. Cottonii and Spinosum Cultivation Handbook. FMC Food
Ingredients Division, Philadelphia, 52 pp.
Ask, E. I., Batibasaga, A., Zertuche-Gonzales, J. A. and de San, M., (in
press). Introducing cultivated varieties of Kappaphycus alvarezii (Doty) to
nonendemic locations: suggested quarantine and introduction procedures plus a
study of the impact of introduction to a Fiji Islands' lagoon. Journal of Applied
Phycology, . 63
Ask, E., Ledua, E., Mario, S., Batibasaga, A., 2001. Developing the cottonii
(Kappaphycus alvarezii) cultivation industry in the Fiji Islands. (abstract only)
XVIIth International Seaweed Symposium: Programme and Abstracts. Cape Town,
South Africa.
Ask, E.I. & Azanza, R.V. 2002. Advances in cultivation technology of commercial
eucheumatoid species: a review with suggestions for future research. Aquaculture
206 pp.257-277.
Atmadja, W. S. & sulistijo, 1980. Experimental cultivation of red algal genera
Eucheuma and Gracilaria in the lagoon of Pari Island, Indonesia. Tropical Ecology &
Development, 1980: 1121-1126.Azanza-Corrales, R., 1990. The farmed
Eucheuma species in Danajon Reef, Philippines: vegetative and reproductive
structures. J. Appl. Phycol. 2, 57-62.
monograph index
Page 75
next
Azanza-Corrales, R., Aliaza, T.T., 1999. In vitro carpospores release and

germination in Kappaphycus alvarezii (Doty) Doty from Tawi-Tawi, Philippines.
Botanica Marina 42, 281-284.
Azanza-Corrales, R., Dawes, C.J., 1989. Wound healing in cultured Eucheuma
alvarezii var. tambalang Doty. Botanica Marina 32, 229-234.
Azanza-Corrales, R., Sa-a, P., 1990. The farmed Eucheuma species
(Gigartinales, Rhodophyta) in Danajon Reef, Philippines: carrageenan properties.
Hydrobiologia 204-205, 521-525.
Azanza-Corrales, R., Mamauag, S.S., Alfiler, E., Orolfo, M.J., 1992.
Reproduction in Eucheuma denticulatum (Burman) Collins and Harvey and
Kappaphycus alvarezii (Doty) Doty farmed in Danajon Reer, Philippines.
Aquaculture 103, 29-34.
Azanza-Corrales, R., Aliaza, T.T., Montailo, N.E., 1996. Recruitment of
Eucheuma and Kappaphycus on a farm in Tawi-Tawi, Philippines. Hydrobiologia 326327, 235-244. Baltazar, G.Q., 1992. Growth rates of Eucheuma cottonii and
Eucheuma spinosum in the occasionally exposed and fully submerged environment.
In: Calumpong, H., Meflez, E. (Eds.), Proceedings of the 2nd RP-USA Phycology
Symposium/workshop. Cebu City, Philippines.
Barbaroux, 0., Perez, R., Dreno, J.P., 1984. L'algue rouge Eucheuma spinosum
possibilities d'exploitation et de culture aux Antilles. Sci. Peche, Bull. Inst. Peches
Marit. 348, 2-9.
Barraca, R.T.,1989. Performance of Eucheuma seaweed in Indonesia. Part 1.
Agronomic characters. Tech. Bull. FMC Marine Colloids Divn. Unpublished. Barraca,
R. T., (1998). Seaweed assessment report (SAR) Malalag Bay Enterprise
Development Zone. Coastal Resource Management Project (CRMP), Cebu City,
Philippines.
Barraca, R. T., (1998). Seaweed assessment report (SAR) Negros Oriental
Enterprise Development Zone. Coastal Resource Management Project (CRMP), Cebu
City, Philippines.
Barraca, R. T., (1998). Seaweed assessment report (SAR) San Vincente Bay
City, Philippines.
Barraca, R. T., (1998). Seaweed assessment report (SAR) Sarangani Bay
City, Philippines.
Barraca, R. T., (1999). Guided Training Manual: Development of seaweed farming
enterprise. Coastal Resource Management Project (CRMP), Cebu City, Philippines.
Bixler, H.J., 1995. Recent developments in manufacturing and marketing
carrageenan. Hydrobiologia 326-327, 35-57.
Blakemore, W.R., 1990. Post harvest treatment and quality control of Eucheuma
seaweeds. In: Adarns, T., Foscarini, R. (Eds.), Proceedings of the Regional
Workshop on Seaweed Culture and Marketing. South Pacific Aquaculture
Development Project, Food and Agriculture Organization of the United Nations,
Suva, Fiji, pp.48-52.
Bodvin, T., 1995. Clean technology in aquaculture; a production without waste
products. Proceedings of the XVth International Seaweed Symposium. Valdivia,
Chile.
SuriaLink 1-0703

Bibliography Bor-Dix
Borgesen, F., 1943. Some marine algae from Mauritius. III. Rhodophyceae. K.
dan. Vidensk. Selsk. Biol. Meddr 19(1): 5-85. Borgesen, F., 1950. Some Marine
Algae from Mauritius. K. dan. Vidensk. Selsk. Biol. Meddr 18(11): 5-44.
Borgesen, F., 1953. Some Marine Algae from Mauritius. K. dan. Vidensk. Selsk.
Biol. Meddr 21(9): 5-62.
Bureau of Fisheries and Aquatic Resources (BFAR), 1974. Illustrated
Instructions for Private Eucheuma Farmers. Bureau of Fisheries and Aquatic
Resources, Manila, Philippines.
Buschmann, A., M. Troell & N. Kautsky, 1995. Integrated tank cultivation of
salmonids and Gracilaria chilensis. Proceedings of the XVth International Seaweed
Symposium. Valdivia, Chile.
Braud, J.P., Perez, R., 1978. Farming on a pilot scale of Eucheuma spinosum
(Florideophyceae) in Djibouti waters. In: Jenson, A., Stein, J. (Eds.), Proceedings of
the 9th International Seaweed Symposium. Science Press, Princeton, pp.533-539.
Braud, J.P., Perez, R., Lacherade, G., 1974. Etude des possibilite's d'adaptation
de l'algue rouge Eucheurna spinosum aux cotes des Afars et des Issas. Sci. Peche,
Bull. Inst. Peches Marit. 238, 1-16, Juillet-nout.
Braud, J.P., R. Perez & G. Lacherade, 1974. Etude des possibilities d' adaptation
de l'algue rouge Eucheuma spinosum aux cotes du Territoire francais des Afars et
des Issas. Sci. Peche, No. 238, 16 pp.
Cheney, D. P., 1986. Genetic engineering in seaweeds: applications and current
status. Nova Hedwegia 81: 22-29. 208. Cheney, D.P., Dawes, C.J., 1981.
Taxonomic studies of the Florida species of Eucheuma (Rhodophyta, Gigartinales):
I. Initial considerations. In: Levring, T. (Ed.), Proceedings of the l0nth International
Seaweed Symposium, vol.8. Walter de Grnyter, Berlin, pp.59-66.
Cheney, D.P., Metz, B., Levine, I., Rudolph, B., 1998. Genetic manipulation and
strain improvement of seaweeds for aquaculture. Book of Abstracts, Aquaculture
'98 Las Vegas, Nevada, p.105.
Collen, J., Mtoletra, M., Abrahamsson, K., Semesi, A. and Pederson, M.,
(1995). Framing and physiology of the red algae Eucheuma: growing commercial
importance in East Africa. Ambio, 24 (7) : 497-501.
Critchley, A. T. and Ohno, M. (eds.), 1998. Seaweed Resources of the World,
Japan International Cooperation Agency, Yokosuka: 343-346.Dawes, C. J., 1979.
Physiological and biochemical comparisons of Eucheuma spp. (Florideophyceae)
yielding iota-carrageenan. Proc. int. Seaweed Symp. 9: 199-207.
Dawes, C.J., 1984. Physiological ecology of Eucheuma species that contain iota
carrageenan from the Pacific and Caribbean Oceans. 1st Philippines-U.S. Phycology
Workshop. Manila, Philippines, October 24-November 5th, 1984, pp.1-8.
Dawes, C.J., 1989. Temperature acclimation in cultured Eucheuma isiforme from
Florida and E. alvarezii from the Philippines. J. AppL PhycoL 1, 59-69. Dawes, C.J.,
1989. Temperature acclimation in cultured Eucheuma isiforme from Florida and E.
alvarezii from the Philippines. J. AppL PhycoL 1, 59-69.
monograph index
Page 76
next
Dawes, C. J., Earle, S.A. & Croley,F.C., 1967. the offshore benthic flora of the
southwest coast of Florida. Bull. Mar. Sci. 17: 211-231.
Dawes, C.J., 1992. Irradiance acclimation of the cultured Philippines seaweeds,
Kappaphycus alvarezii and Eucheuma denticulatum. Botanica Marina 35, 189-195.
Dawes, C.J., Koch, E.W., 1991. Branch, micropropagule and tissue culture of the
red algae Eucheuma denticulatum and Kappaphycus alvarezii farmed in the
Philippines. J. Appl. Phycol. 3, 247-257.
Dawes, C.J., A.C. Mathieson & D.P. Cheney, 1974. Ecological studies of
Floridian Eucheuma (Rhodophyta, Gigartinales). I. Seasonal growth and
reproduction. Bull. Mar. Sci. 24: 235-273.
Dawes, C.J., Trono, G.C., Lluisma, A.O., 1993. Clonal propagation of Eucheuma
denticulatum and Kappaphycus alvarezii for Philippine seaweed farms.
Hydrobiologia 260-261, 379-383.
Dawes, C.J., Lluisma, A.O., Trono, G.C., 1994. Laboratory and field growth
studies of commercial strains of Eucheuma denticulatum and Kappaphycus alvarezii
in the Philippines. 3. Appl. Phycol. 6, 21-24. Dawson, E.Y., 1961. Marine Red
Algae of pacific Mexico. Part 4. Gigartinales. Pacif. Nat. 2: 191-343.
Delmendo, M. N., Alvarez, V. and Rabanal, H. R., (1992). The evolution of
seaweed farming development and its relevance to rural agro-industrial
development of coastal communities in the Philippines. Department of Fisheries,
Bureau of Fisheries and Aquatic Resources, Manila, Philippines. 86
De Bruin, G.H.P, Russell, B.C., Bogusch, A., 1994. The marine fishery resources
of Sri Lanka, FAO Species Identification Field Guide for Fishery Purposes, Food and
Agriculture Organization of the United Nations, Rome, ISBN 92-5-103293-9.
De Paula, E.J., Pereira, R.T.L., Ostini, S., 1998. Intreducao de especies exoticas
de Eucheuma e Kappaphycus (Gigartinales, Rhodophyta) para fms de maricultura
no litoral Brasileiro: abordagem te6rica e experimental. In: de Paula, E.J., CordeiroMarino, M., Pupo Santos, D., Fujii, M., Plastino, E.M., Yokoya, N. Eds.), W
Congresso Latino Americano de Ficologia, II Reuniao Ibero-Americana de Ficologia
eVil Reuniao Brasileira de Ficologia, Sao Paulo, Brazil, pp.340-357.
De Paula, E.J., Pereira, R.T.L., Olmo, M., 1999. Strain selection in Kappaphycus
alvarezii var. alvarezii (Doty) Doty ex P Silva (Rhodophyta, Solieriaceae) using
tetraspore progeny. 3. AppI. Phycol. 11(1), 111 - 121.
De Reviers, B., 1989. Realisation d'Une Ferme de Culture Industrielle de
Eucheuma aux Maldives. Oceanis 15 (5), pp. 749-752.
Dixon, P.S., 1962. Taxonomic and nomenclature notes on the Florideae, III. Bot.
Notiser. 115: pp. 245-260.Doty, J.E. & M.S. Doty, 1973. Abrasion in the
measurement of water motion with the cold-card technique. Bull. South. Calif.
Acad. Sci. 72: 40-41.
SuriaLink 1-0703

Bibliography Dot-Gle
Doty, M. S., 1971. Physical factors in the production of tropical benthic marine
algae. In J.D. Costlow (ed.), Fertility of the Sea. Vol. 1. Gordon & Breach Science
Publishers, New York. pp. 99-121.
Doty, M.S., 1973. Farming the red seaweed, Eucheuma, for Carrageenans.
Micronesia 9 (1), 59-73.
Doty, M.S., 1978a. Eucheuma-current marine agronomy. In: Krauss, R.W. (Ed.),
The Marine Plant Biomass of the Pacific Northwest Coast. Oregon State Univ. Press,
Corvallis, OR, pp.203-214.
Doty, M.S., 1978b. Status of marine agronomy with special reference to the
tropics. In: Jensen, A., Stein, JR. (Eds.), Proceedings of the 9th International
Seaweed Symposium. Science Press, Princeton, pp.34-58.
Doty, M.S., 1980. Outplanting Eucheuma species and Gracilaria species in the
Tropics. In: Abbott, IA., Foster; Doty, M.S., 1985. Eucheuma alvarezii sp. nov.
(Gigartinales, Rhodophyta) from Malaysia. In I.A. Abbott & J.N. Norris (eds),
Taxonomy of Economic Seaweeds. California Sea Grant College Program, Univ. of
California, La Jolla, California, pp. 37-45.
Doty, M. S. 1982. Realizing a Nation's Potential in Phycology. In R.T. Tsuda & Y-M.
Chiang (eds), Proceedings of the Republic of China---United States Cooperative
Science Seminar on Cultivation and Utilization of Economic Algae. University of
Guam Marine Laboratory, Mangilao, Guam, pp. 1-7.
Doty, M. S., 1987. The production and use of Eucheuma. In M.S. Doty, J.F. Caddy
& B. Santelices (eds), Case Studies of Seven Commercial Seaweed Resources. FAO
Fisheries Technical Paper 281, Food and Agriculture Organization of the United
Nations, Rome, pp. 123-164.
Doty, M.S., 1985a. Eucheuma alvarezii, sp. Nov. (Gigartinales, Rhodophyta) from
Malaysia. In: Abbott, IA, Norris, J.N. Eds.), Taxonomy of Economic Seaweeds: With
Reference to Some Pacific and Caribbean Species. California Sea Grant College
Program. Rep. T-CSGCP-01 1, La Jolla, California, pp.37-45.
Doty, M.S., 1985b. Eucheuma species (Solieriaceae, Rhodophyta) that are major
sources of carrageenan. In: Abbott, LA., Norris, J.N. Eds.), Taxonomy of Economic
Seaweeds: With Reference to Some Pacific and Caribbean Species. California Sea
Grant College Program. Rep. T-CSGCP-0l 1, La Jolla, California, pp. 47-61. Doty,
M.S., 1987. The production and use of Eucheuma. In: Doty, MA., Caddy, J.F.,
Santelices, B. (Eds.), Case Studies of Seven Commercial Seaweed Resources. FAO
Fish. Tech. Pap., 281 Rome, pp. 123-161. Doty, M.S., 1988. Prodomus ad
systematica Eucheumatoideorum: a tribe of commercial seaweed related to
Eucheuma (Solierieaceae, Gigartinales). In: Abbott, LA. (Ed.), Taxonomy of
Economic Seaweeds: With Reference to Some Pacific and Caribbean Species,
Volume II. California Sea Grant College Program. Rep. T-CSGCP~l 1, La Jolla,
California, pp.159-207.
Doty, M.S., Alvarez, V.B., 1973. Seaweed farms: a new approach for U.S.
industry. Proceedings of the 9th Annual Conf. Proceedings, University of Hawaii,
Hawaii, pp.701-708.
Doty, M.S., Alvarez, V.B., 1975. Status, problems, advances and economics of
Eucheuma farms. Mar. Technol. Soc. 3.9, 30-35.
Doty, M.S., Alvarez, V.B., 1981. Eucheuma farm productivity. In: Fogg, G.E.,
Jones,W.E., Eds.), Proceedings of the Eight International Seaweed Symposium. The
Marine Science Laboratory, Menai Bridge, Hawaii, pp. 688-691.
monograph index
Page 77
next
Doty, M. S., B. J. Cook, J. R. fisher, I. A. Levine & E. K. Zablackis, 1986.

Experiments with Gracilaria in Hawaii 1983-1985. Hawaii Bot. Sci. Pap. #46, Univ.
of Hawaii, Honolulu, 486 pp.
Doty, M.S. & J.N Norris, 1985. Eucheuma species (Solieriaceae, Rhodophyta)
that are major sources of carrageenan. In I.A. Abbott & J.N. Norris (eds),
Taxonomy of Economic Seaweeds. California Sea Grant College Program Report No.
T-CSGCP-011, 47-61.
Doty, M.S. & G.A. Santos, 1978. Carrageenans from tetrasporic and cystocarpic
Eucheuma species. Aquat. Bot. 4: 143-149.
Epiard-Lahaye, M., 1988. Effects of ammonium, nitrate and phosphate on the
growth of Cystoseira stricta (Phaeophyta, Fucales) cuttings in culture. Cryptogam.
Algol. 9, pp. 211-239.
Fa'anunu, U., 1990. Tonga. In: Adams, T., Foscarini, R. (Eds.), Proceedings of the
Regional Workshop on Seaweed Culture and Marketing. South Pacific Aquaculture
Suva, Fiji, pp.25-31.
Fang, T.C., 1983. A summary of the genetic strains of Laminaria japonica in China.
In C.K. Tseng (ed.), Proceedings of the Joint China-U.S. Phycology Symposium.
Science Press, Beijing, China, pp. 123-136.
Fang, T. C., C. Y. Wu, B. Y. Jiang, J. J. Li & K.Z. Ren, 1963. The breeding of a
new variety of Haidai (Laminaria japonica). Sci. Sin. 12, pp. 1011-1018. Fletcher,
R.L., 1995. Epiphytism and fouling in Gracilaria cultivation: an overview. J. Appl.
Phycol. 7, pp. 325-333.
Gabrielson, P.W., 1983. Vegetative and reproductive morphology of Eucheuma
isiforme (Solieriaceae, Gigartinales, Rhodophyta). J. Phycol. 19, pp. 45-52.
Gabrielson, P. W. & D. P. Cheney, 1987. Morphology and taxonomy of
Meristiella Gen. Nov. (Solieriaceae, Rhodophyta). J. Phycol. 23, pp. 481-493.
Gabrielson, P.W. & G.T. Kraft, 1984. The marine algae of Lord Howe Island
(N.S.W.): the Family Solieriaceae (Gigartinales, Rhodophyta). Brunonia 7, pp. 217251.
GEM, 1999. Filipinos Test New Methods. Fish Farming International, March 1999,
Zamboanga City, Philippines, pp.26-27, March.
Gentle, T., 1990. Tuvalu. In: Adams, T., Foscarini, R. (Eds.), Proceedings of the
Suva, Fiji, 14-17 November, 1989, pp.32-33.
Glenn, E.P. & M.S. Doty, 1981. Photosynthesis and respiration of the tropical red
seaweeds, Eucheuma striatum (Tambalang and Elkhorn varieties) and E.
denticulatum. Aquat. Bot. 10, pp. 353-364.
Glenn, E.P., Doty, M.S., 1990. Growth of the seaweeds Kappaphycus alvarezii, K
striatum and Eucheuma denticulatum as affected by environment in Hawaii.
Aquaculture 84, pp. 245-255.
Glenn, E.P., Doty, M.S., 1992. Water motion affects the growth rates of
Kappaphycus alvarezii and related red seaweeds. Aquaculture 108,pp.233-246.
Glenn, E.P. & M.S. Doty, 1981. Photosynthesis and respiration of the tropical red
seaweeds, Eucheuma striatum (Tambalang and Elkhorn varieties) and E.
denticulatum. Aquat. Bot. 10: 353-364.
SuriaLink 1-0703

Bibliography Gom-Liu
Gomez, E.D., Azanza-Corrales, R., 1988. Polyculture of seaweeds with marine
animals. Report on the Training Course on Seaweed Farming. Manila Philippines, 221 May. FAO, pp.81-85.
Gomez, E. D., Buieb, R. A. and Arc, E., 1983. Studies on the predators of
commercially important seaweeds. Fish. Res. J. Philippines, 8 : 1-17.
Gordon, E. M. & E. L. McCandless, 1973. Ultrastructure and histochemistry of
Chondrus crispus Stackhouse. Proc. N. S. Inst. Sci. 27 (suppl. 1973): 111-133.
Greer, C. W. & W. Yaphe, 1984. Characterization of hybrid (beta-kappa-gamma)
carrageenan from Eucheuma gelatinae J. Agardh (Rhodophyta, Solieriaceae) using
carrageenases, infrared and 13C-nuclear magnetic resonance spectroscopy.
Botanica Marina 27, pp. 473-478.Grigg, R.W., 1988. Paleoceanography of coral
reefs in the Hawaiian-Emperor Chain. Science 240, pp. 1737-1743.
Hanisak, M.D., 1987. Cultivation of Gracilaria and other macroalgae in Florida for
energy production. In K.T. Bird & P. H. Benson (eds), Seaweed Cultivation for
Renewable Resources. Elsevier, Amsterdam, pp. 191-218.
Harlin, M. M., 1978. Nitrate uptake by Enteromorpha spp. (Chlorophyceae):
applications to aquaculture systems. Aquaculture 15: 373-376.
Harvey, W.H., 1853. Nereis Boreali-Americana,... II: Rhodospermeae. [First
Issue]. John van voorst, Washington-London. ii + 258 pp., pls 13-36.
Hindley, F., (1999). The environmental impacts of seaweed farming (genus
Eucheuma) in the Philippines with a particular reference to seagrass. In: Imperial
College of Science, Technology and MedicineUniversity of London, London. 69
Horstman, M.A., A. Colina & W. Shramin, 1977. some aspects of the culture of
Eucheuma. Mar. Res. Indonesia 1977 (17), p. 145.
Hurtado-Ponce, A.Q., 1992. Cage culture of Kappaphycus alvarezii var.
tambalang (Gigartinales, Rhodophyceae). J. Appl. Phycol. 4, pp. 311-313.
Hurtado-Ponce, A., 1995. Polyculture of Gracilaria heterocladia (Zhang et Xia)
and Penaeus monodon Fabricius in brackishwater ponds. Proceedings of the XVth
International Seaweed Symposium. Valdivia, Chile.
Jimenez del Rio, M., Z. Ramazanov & G. Garcia-Reina., 1995. Ulva rigida
biofilters for reduction of dissolved inorganic nitrogen in fish farm effluents.
Proceedings of the XVth International Seaweed Symposium. Valdivia, Chile.
Johnstone, R. W. and Olafsson, E., (1995). Some environmental aspects of
open water algal cultivation: Zanzibar, Tanzania. Ambio, 24 (7-8) : 465-469.
Juanich, G.L., 1988. Manual on Seaweed Farming 1. Eucheuma sp.,
ASEAN/SF/88/Manual No.2 ASEANI UNDP/FAO Regional Small-Scale Coastal
Fisheries Development Project, Manila, Philippines.
Kapraun, D.F., Lopez-Bautista, J., 1997. Karyology, nuclear genome
quantification and characterization of the carrageenophytes Eucheuma and
Kappaphycus (Gigartinales). J. Appl. Phycol. 8, pp. 465-471.
Kawashima, S., 1984. Kombu culture in Japan for human foodstuff. Jap. J. Phycol.
32, pp. 379-394.
Kraft, G.T., 1969. Eucheuma procrusteanum, a new red algal species from the
Philippines. Phycologia 8, pp. 215-219.
monograph index
Page 78
next
Kraft, G.T., 1972. Preliminary studies of Philippine Eucheuma species

(Rhodophyta). part 1, taxonomy and ecology of Eucheuma arnoldii. Pacif. Sci. 26,
pp. 318-335.
Kylin, H., 1932. Die Florideenordnung Gigartinales. Lunds Univ. Arsskr., N.F. Avd.
2, 28 (8), pp. 1-88.
Kylin, H., 1956. Die Gattungen der Rhodophyceen. C.W.K. Gleerups, Lund, 673
pp.
Largo, D.B., Fukami, K., Nishijima, T., 1995a. Occasional pathogenic bacteria
promoting ice-ice disease in the carrageenan-producing red algae Kappaphycus
alvarezii and Eucheuma denticulatum (Solieriaceae, Gigartinales, Rhedophyta). 3.
Appl. Phycol. 7, pp. 545-554.
Largo, D.B., Fukami, F., Nishijima, T., Olino, M., 1995b. Laboratory-induced
development of the ice-ice disease of the farmed red algae Kappaphycus alvarezii
and Eucheuma denticulatum (Solieriaceae, Gigartinales, Rhodophyta). J. Appl.
Phycol. 7, pp. 539-543.
Largo, D. B., Fukami, K., Adachi, M. and Nishijima, T., (1998).
Immunofluorescent detection of ice-ice disease-promoting bacterial strain Vibrio sp.
P11 of the farmed macroalga, Kappaphycus alvarezii (Gigartinales, Rhodophyta).
Journal of Marine Biotechnology, 6 (3) : 178-182.
Largo, D. B., Fukami, K. and Nishijima, T., (1999). Time-dependent
attachment mechanism of bacterial pathogen during ice-ice infection in
Kappaphycus alvarezii (Gigartinales, Rhodophyta). Journal of Applied Phycology, 11
(1) : 129-136.
Lawson, C.J., D. A. Rees, D.J. Stanley & N.F. Stanley, 1973. Carrageenans.
Part VIII. Repeating structures of galactan sulphates from Furcellaria fastigiata,
Gigartina canaliculata, Gigartina chamissoi, Gigartina atropurpurea, Ahnfeltia
durvillaei, Gymnogongrus furcellatus, Eucheuma cottonii, Eucheuma spinosum,
Eucheuma isiforme, Eucheuma uncinatum, Agardhiella tenera, Pachymenia
hymantophora and Gloipeltis cervicornis. J. Chem. Soc. Perkin. I 1073, pp. 21772182.
Lignell, A. & M. Pedersen, 1987. Nitrogen metabolism in Gracilaria secundata,
Harv. Hydrobiologia 151/152, pp. 431-441.
Li, R., Li, 3., Wu, C.Y, 1990. Effect of ammonium on growth and carrageenan
content in Kappaphycus alvarezii (Gigartinales, Rhodophyta). Hydrobiologia pp. 204205, 499-503.
Lim, J.R., 1982. Farming the Ocean (The Genu Story). Historical Conservation
Society, Manila, No. 36, pp. 1-143. (With appendices).
Lim, JR., Porse, H., 1981. Breakthrough in the commercial culture of Eucheuma
spinosum in Northern Bohol, Philippines. In: Levring, T. Ed.), Proceedings of the
10th International Seaweed Symposium. Walter de Grayter, Berlin, pp.601-606.
Lirasan, T., Twide, P., 1993. Farming Eucheuma in Zanzibar, Tanzania.
Hydrobiologia 260-261, pp. 353-355.
Liu, S. & P. Zhuang, 1984. The commercial cultivation of Eucheuma in China.
Proc. int. Seaweed Symp. 11, pp. 243-245.
SuriaLink 1-0703

Bibliography Lux-Ola
Luxton, D.M., Luxton, PM., 1999. Development of commercial Kappaphycus
production in the Line Islands, Central Pacific. Hydrobiologica 398/399, pp. 477486.
Luxton, D. M., 1993. Aspects of the farming and processing of Kappaphucus and
Eucheuma in Indonesia. Hydrobiologia, 260/261 : 365-371.
Luxton, L.M., Robertson, M., Kindley, M.J., 1987. Farming of Eucheuma in the
South Pacific Islands of Fiji. Hydrobiologia 151-152, pp. 359-362. Manzano, J.V.,
Manzano, V.B., 1992. Development of culture techniques for local abalone
integrated to Eucheuma farm (Abstract). In: Proceedings of the 2nd RP-USA
Phycology Symposium-Workshop, Cebu City, Philippines, p.257. Mairh, OP., SoeHtun, U., Ohno, M., 1986. Culture of Eucheuma striatum (Rhodophyta,
Solieriaceae) in subtropical waters of Shikoku, Japan. Botanica Marina 29, pp. 185191.
Mairh, O. P. & A. Tewari, 1994. Studies on a new asexual propagule of
Kappaphycus striatum (Soleiriacea, Rhodophyta). Phycologia 33 (1), 62-64.
Mairh, O. P., S. T. Zodape, A. Tewari & M. R. Rajyaguru, 1995. Culture of
Marine Alga Kappaphycus striatum (Schmitz) Doty on the Suarashtra region, west
coast of India. Indian Journal of Marine Sciences 24, March 1995 pp. 24-31.
Mathieson, A.C., Dawes, C.J., 1974. Ecological studies of Floridian Eucheuma
(Rhodophyta, Gigartinales): 11. Photosynthesis and respiration. Bull. Mar. Sci. 34,
pp. 274-285.
McCandless, E.L., J.S. Craigie & J.A. Walter, 1973. Carrageenans in the
gametophytic and sporophytic stages of Chondrus crispus. Planta 112, pp. 201-212.
Mollion, J., Braud, J.P., 1993. A Eucheuma (Solieriaceae, Rhodophyta) cultivation
test on the south-west coast of Madagascar. Hydrobiologia 260-261, pp. 373-378.
Moon, R.E. & C.J. Dawes, 1976. Pigment changes and photosynthetic rates under
selected wavelengths in the growing tips of Eucheuma isiforme (C. Agardh var.
denudatum Cheney ) during vegetative growth. Br. Phycol. J. 11, pp. 165-174.
Mshigeni, K.E., 1982. Seaweed resources in Tanzania: a survey of potential
sources for industrial phycocolloids and for other uses. Marine Algae in
Pharmaceutical Science. Vol. 2. Walter de Gruyter, Berl., pp. 131-174.
Mshigeni, K.E. & A.K. Semesi, 1977. Studies on carrageenans from the economic
red algal genus Eucheuma in Tanzania. Botanica Marina 20, pp. 239-242.
Mshigeni, K. E., (1989). Seaweed faming in tropical seas: the case of Eucheuma
in the western Indian Ocean region. In: Cultivation of Seaweeds in Latin America,
de Oliveira, E. C. and Kautsky, N. (eds.), S. Sebastiao, Sao Paulo: 389-397.
Msuya, F., Ngoile, M. A. and Shunula, J. P., (1997). The impact of seaweed
farming on the macrobenthos of the east coast of Unguja Island, Zanzibar,
Tanzania. University of Dar es Salaam, Zanzibar.
Mtolera, M.S.P., Collen, J., Pedersen, M., Ekdahl, A., Abrahanisson, K.,
monograph index
Page 79
next
Semesi, A.K., 1996. Stress-induced production of volatile halogenated organic

compounds in Eucheuma denticulatum (Rhodophyta) caused by elevated pH and
high light intensities. Eur. J. Phycol. 3, pp. 89-95.
Neish, A.C. & P.F. Shacklock, 1971. Greenhouse experiments (1971) on the
propagation of Irish moss. National Research Council of Canada, Atlantic Regional
Laboratory, Halifax. N.S. Tech. Reg. 14, 25 pp.
Neish, IC., Ask, E.I, 1995. Expert systems in sustainable integrated mariculture.
In: Sustainable Aquaculture 1995 Proceedings, Pacific Congress on Marine Science
and Technology, PACON International, Hawaii, pp. 271-280.
Neushul, M. & B. W. W. Harger, 1987. Nearshore kelp cultivation, yield and
genetics. In K. T. Bird & P. H. Benson (eds), Seaweed Cultivation for Renewable
Resources. Elsevier, Amsterdam, pp. 69-93.
Nicholl, D.S.T., 1996. An Introduction to Genetic Engineering. Cambridge Univ.
Press, Cambridge, 133 pp.
Njoman, I., S. Nuitja, D. Soedharma, M. Ohno & C.A. Orosco, 1987. Studies
on the Culture of Eucheuma cottonii. Nori & Sea Vegetables No. 29, pp. 28-34
Norris, J.N. 1985. Observations on Eucheuma J. Agardh (Solieriaceae,
Rhodophyta), from the Gulf of California Mexico. In I.A. Abbott & J. N. Norris (eds),
Taxonomy of Economic Seaweeds. California Sea Grant College Report No. T-CSGCP011, pp. 63-65.
North, W.J., 1987. Oceanic farming of Macrocystis, the problems and nonproblems. In K.T. Bird & P.H. Benson (eds), Seaweed Cultivation for Renewable
Resources. Elsevier, Amsterdam, pp. 39-67.
Ohmi, H. & I. Shinmura, 1976. Growth of Eucheuma amakusaensis in the field
culture. Bull. jap. Soc. Phycol. 24, pp. 98-102.
Ohno, M., Nang, H.O., Dinh, N.H., Triet, VD., 1995. On the growth of cultivated
Kappaphycus alvarezii in Vietnam. Jpn. J. Phycol. 43, pp. 19-22.
Ohno, M., Nang, H.O., Hirase, 5., 1996. Cultivation and carrageenan yield and
quality of Kappaphycus alvarezii in the waters of Vietnam. J. Appl. Phycol. 8, pp.
431-437.
Ohno, M. & C.A. Orosco, 1987. Growth rate of three species of Eucheuma,
commercial red algae from the Philippines. In I. Umezaki (ed.), Scientific Survey of
Marine algae and their Resources in the Philippine Islands. Privately published by
the Laboratory of Fishery Resources, Graduate School of Agriculture, Kyoto
University, pp. 77-81.
Ohno, M., Largo, D.B., Ikurnoto, T., 1994. Growth rate, carrageenan yield and
gel properties of culture kappa carrageenan producing red alga Kappaphycus
alvarezii (Doty) Doty in the subtropical waters of Shikoku, Japan. J. AppL Phycol. 6,
pp. 1-5.
Okamura, K., 1906. Icones of Japanese Algae. Vol. II, #41, Plates LXI & LXII.
Privately published by the author.
Okamura, K., 1936. Nippon Kaiso-shi. [Japanese Algae]. Uchidarokakuho, Tokyo.
964 pp.
Okuda, T., Neushul, M., 1983. Sedimentation studies of red algal spores. J.
Phycol. 17, pp. 113-118.
Olafsson, E., Johnstone Ron, W. and Ndaro Simon, G. M., (1995). Effects of
intensive seaweed farming on the meiobenthos in a tropical lagoon. Journal of
Experimental Marine Biology & Ecology, 191 (1) : 101-117.
SuriaLink 1-0703

Bibliography Pad-Sid
Padilla, J.E., Lampe, H.C., 1989. The economics of seaweed farming in the
Philippines. NAGA, ICLARM Q., 12 (3), pp. 3-5.
Parker, H.S., 1974. The culture of the red algal genus Eucheuma in the
Philippines. Aquaculture 3, pp. 425-439.
Pedersen, M., Collen, J., Abrahamsson, K., Mtolera, M., Semesi, A., GarciaReina, G., 1996. The ice-ice disease and oxidative stress of marine algae. In:
Bjork, M., Semesi, A.K., Pedersen, M., Bergman, B. Eds.), Current Trends in Marine
Botanical Research in the East African Region. Proceeding of the 3-10 December
1995 Symposium on the Biology of Microalgae, Macroalgae and Seagrasses in the
Western Indian Ocean. Swedish International Development Agency (SIDA),
Stockholm, Sweden, pp.11-24.
Penman, A. & D.A. Rees, 1973. Carrageenans. Par. IX Methylation analysis of
galactan sulphates from Furcellaria fastigiata, Gigartina canaliculata, Gigartina
chamissoi, Gigartina atropurpurea, Eucheuma isiforme, Eucheuma uncinatum
Agardhiella tenera, Pachymenia hymantophora and Gloipeltis cervicornis. Structure
of E-carrageenan (epsilon-carrageenan). J. Chem. Soc. Perkin I. 1973, pp. 21832187.
Perez, R., 1992. Essais de cultures hors de la zone asiatique. In: Perez, R. (Ed.),
La Culture Des Algues Marines Dans le Monde. IFREMER Centre de Brest, Plouzane,
France, pp.213-215.
Perez, R., Braud, IP., 1978. Possiblite' d'Une Culture Industielle de L'Algau Rouge
Eucheuma spinosum Dans Le Golfe de Tadjourah. Sci. Peche, Bull. Inst. Peches
Marit 285, pp. 1-27.
Petrell, R., 1995. Integrated culture of salmonids and seaweeds in open systems.
Proceedings of the XVth International Seaweed Symposium. Valdivia, Chile.
Polne-Fuller, M. & A. Gibor, 1987. Tissue Culture of Seaweeds. In K. T. Bird & P.
H. Benson (eds), Seaweed cultivation for renewable resources. Elsevier,
Amsterdam, pp. 219-239.
Prakash, J., 1990. Fiji. In: Adams, T., Foscarini, R. (Eds.), Proceedings of the
Qian, PY, Wu, C.Y, Wu, M., Xie, YK., 1996. Integrated cultivation of the red alga
Kappaphycus alvarezii and the pearl oyster Pinctada martensi. Aquaculture 147, pp.
21-35.
Reddy, C. R. K. Raja Krishna Kumar, G., Siddhanta, A. K. & Tewari, A. 2003.
In vitro somatic embryogenesis and regeneration of somatic embryos from
pigmented callus of Kappaphycus alvarezii (Doty) Doty (Rhodophyta, Gigartinales).
J. Phycol. 39, pp. 610-616.
Ricohermoso, M.A., Deveau, L.E., 1979. Review of commercial propagation of
Eucheuma (Florideophyceae) clones in the Philippines. In: Jensen, A., Stein, IR.
(Eds.), Proceedings of the 9th International Seaweed Symposium. Science Press,
Princeton, pp.525-531.
Rincones, R.E., Rubio, J.N., 1999. Introduction and commercial cultivation of the
red alga Eucheuma in Venezuela for the production of phycocolloids. World
Aquacult. Mag. 30 (2), pp. 57-61.
monograph index
Page 80
next
Robertson, M., 1989. Growing seaweed in Fiji. In: Adams, T., Foscarini, R. (Eds.),
Proc. of the Regional Workshop on Seaweed Culture and Marketing. South Pacific
Aquaculture Development Project, Food and Agriculture Organization of the U.N.,
Robledo, D., (1998). The Seaweed Resources of Mexico. In: Seaweed Resources
of the World, Critchley, A. T. and Ohno, M. (eds.), Japan International Cooperation
Agency, Yokosuka: 331-342.
Rodgers, S.K., Cox, E.F., 1999. Rate of spread of introduced Rhodophytes
Kappaphycus alvarezii (Doty), Kappaphycus striatum and Gracilaria salicornia and
their current distributions in Kane'ohe Bay, O'ahu, Hawaii. Pac. Sci. 53 (3), pp. 232241.
Ronquillo, IA., Gabral-Llana, M.E., 1989. The Philippines' Eucheuma seaweed
industry. Fish. Res. J. Philipp. 14, pp. 23-29. Russell, D.J., 1982. Introduction of
Eucheuma to Fanning Atoll, Kiribati, for the purpose of Mariculture. Micronesia 18
(2), pp. 35-44.
Russell, D. J., 1983. Ecology of the imported red seaweed Eucheuma striatum
Schmitz on Coconut Island, Oahu, Hawaii. Pac. Sci. 37, pp. 87-108.
Russell, D. J., (1987). Introduction and establishment of alien marine algae.
Bulletin of Marine Science, 41 (2) : 641-642.
Russell, D. J., (1992). The ecological invasion of Hawaiian reefs by two marine
red algae, Acanthophora spicifera (Vahl) Boerg. and Hypnea musciformis (Wulfen)
J. Ag., and their association with two native species, Laurencia nidifica J. Ag. and
Hypnea cervicornis J. Ag. ICES Marine Science Symposia, 194 : 12-13.
Santos, G.A., 1989. Carrageenans of species Eucheuma J. Agardh Kappaphycus
Doty (Solieriaceae, Rhodophyta). Aquatic Botany 36, pp. 55-67.
Schmitz, O.C., 1985. Marine Florideen von Deutsch Ost-Afrika. Bot. Jahrb. Syst.
21, pp. 298-544.
Schramm, W., Dy, D.T., Gualberto, E.P., Orasco, C.A., Yap, TN., 1984.
Aspects of nutrient dynamics in a Eucheuma farm. First Philippines-U.S. Phycology
Workshop. University of the Philippines, Quezon City, Philippines, pp. 1-3.
Senayabu, Y. & Y. Loya, 1977. Seasonal occurrence of benthic-algae
communities and grazing regulation by sea urchins as the coral reefs of Eilat, Red
Sea. Proc. Third Int. Coral Reef Symp., pp. 383-389.
Serpa-Madrigal, A., Areces, A.J., Cano, M., Bustamante, G., 1997.
Depredacion Sobre Las Carragenofitas Comerciales Kappaphycus alvarezii (Doty)
Doty y K. striatum (Schmitz) Doty (Rhodophyta: Gigartinales) Introducidas en
Cuba. Rev. Invest. Mar.18 (1), pp. 65-69.
Setchell, W. A. & N. L. Gardner, 1924. New marine algae from the Gulf of
California. Proc. Calif. Acad. Sci. 4th Ser. 12, pp. 695-949.
Shinmura, I., 1975. Some observations on Eucheuma amakusaensis. Bull. Jap.
Soc. Phycol. 23: 47-52.
SID-MAFF, 1988. Statistics and Information Department, Ministry of Agriculture,
Forestry and Fisheries, Government of Japan, 1988. Fisheries Statistics of Japan.
Assn. Agriculture and Forestry Statistics, Tokyo, 64 pp.
SuriaLink 1-0703

Bibliography Smi-Vre
Smith, M.T., 1990. Solomon Islands. In: Adams, T., Foscarini, R. (Eds.),
Proceedings of the Regional Workshop on Seaweed Culture and Marketing. South
Pacific Aquaculture Development Project, Food and Agriculture Organization of the
United Nations, Suva, Fiji, 14-17 November, 1989, pp.21-24.
Soegiarto, A. & Sulistijo, 1986. The potential of marine algae for biotechnological
products in Indonesia. In, Workshop on Marine Algae Biotechnology. National
Academy Press, Washington, D. C., pp. 3-15.
Soetjodinoto, 1969. Is the cultivation of seaweed (Eucheuma spinosum and
Eucheuma edule) in Indonesia technically possible and economically justified?
IPFC/C68/TECH 21 at 13th Session, IPFC, Brisbane, Australia, October, 1968, 4 pp.
Solis-Duran, E., Inocencio, D., 1990. Polyculture of giant clams, abalone and
Eucheuma at Bantayan Reef, Dumaguete City. Programs and Abstracts, 1st National
Symposium in Marine Science, UP Marine Science Institute Bolinno Marine
Laboratory, Pangasinan, Philippines, 16-18 May.
Stokoe, P.K. and A.G. Gray, 1990. AQUASITE: a computer site assessment
system for marine coastal aquaculture. Bull. Aquacul. Assoc. Canada 90-4: 94-96.
Sverdrup, H.U., M.W. Johnson & R.H. Fleming, 1942. The Oceans/Their
Physics, Chemistry, and General Biology. Prentice Hall., Englewood Cliffs. N.J.:
1087 pp.
Tanaka, H., 1990. Foreword. In: Adams, T., Foscarini, R. ~ds.), Proceedings of
the Regional Workshop on Seaweed Culture and Marketing. South Pacific
Aquaculture Development Project, Food and Agriculture Organization of the United
Nations, Suva, Fiji, 14-17 November, 1989, pp. ui-iv.
Taniera, T., Tabee, T. and Tebano, T., (1994). Socio-economic of Eucheuma
seaweed farming in Kiribati. University of the South Pacific, Tarawa, Kiribati.
Troell, M., Halling, C., Nilsson, A., Buschmann, A. H., Kautsky, N. and
Kautsky, L., (1997). Integrated marine cultivation of Gracilaria chilensis
(Gracilariales, Bangiophyceae) and salmon cages for reduced environmental impact
and increased economic output. Aquaculture, 156 : 45-61.
Trono, G.C., 1989. Lessons from the history of the seaweed culture in the
Philippines and the trend of seaweed farming in Southeast Asia. In: Adams, T.,
Foscarini, R. (Eds.), Proceedings of the Regional Workshop on Seaweed Culture and
Marketing. South Pacific Aquaculture Development Project, Food and Agriculture
Organization of the United Nations, Suva, Fiji, 14-17 November, 1989, pp.42-47.
Trono, G.C., 1990. A review of the production technologies of tropical species of
economic seaweeds. Regional Workshop on the Culture and Utilization of Seaweeds.
Sponsored by UNDP/FAO and NACA, August 27-September 1, 1990, Cebu City,
Philippines, pp. 1-28.
Trono, G.C., 1992. Eucheuma and Kappaphycus: taxonomy and cultivation. Bull.
Mar. Sci. Fish., Kochi Univ. 12, pp. 51-65. Trono, G.C., 1994. The mariculture of
seaweeds in the tropical Asia-Pacific Region. In: Phang, S.M., Lee, YK., Borowitzka,
M.A., Whitton, B.A. (Eds.), Algal Biotechnology in the Asia-Pacific Region. University
of Malaya, Malaysia, pp.198-210.
monograph index
Page 81
next
Trono, G. C., 1993. Environmental effects of seaweed farming. SICEN Newsletter,

4 (1):1.
Trono, G.C., 1997a. Field Guide and Atlas of the Seaweed Resources of the
Philippines. Bookmark, Makati City, Philippines, 291 pp.
Trono, G. C., 1997b. Eucheuma and Kappaphycus: Taxonomy and cultivation. In:
Seaweed Cultivation and Marine Ranching, Ohno, M. and Critchley, A. T. (eds.),
Japan International Cooperation Agency, Yokosuka: 75-88.
Trono, G.C., Lluisma, A.O., 1992. Differences in biomass production and
carrageenan yields among four strains of farmed carrageenophytes in Northern
Bohol, Philippines. Hydrobiologia 247, 223-227.
Trono, G.C., Ohno, M., 1989. Seasonality in the biomass production of the
Eucheuma strains in Northern Bohol, Philippines. In: Umezaki, 1. Ed.), Scientific
Survey of Marine Algae and their Resources in the Philippine Islands. Monbushio
International Scientific Research Program, Japan, pp.71-80.
Tseng, C.K., 1981. Commercial cultivation. In C.S. Lobban & M.J. Wynne (eds),
The Biology of Seaweeds. Univ. of Calif. Press, Berkeley, pp. 680-725.
Tseng, C.K., 1987. Some remarks on the kelp cultivation industry of China. In K.T.
Birs & P.H. Benson (eds), Seaweed Cultivation for Renewable Resources. Elsevier,
Amsterdam, pp. 147-153.
Uan, J., (1990). Kiribati. In: Proceedings of the Regional Workshop on Seaweed
culture and Marketing. South Pacific Aquaculture Development Project, Adams, T.
and Foscarini, R. (eds.), FAO, Rome: 10-15.
Uy, W.H., Azanza, R.V., Martinez-Goss, M., Israel, A., 1998. Kappaphycus fish
interaction studies. XVIth International Seaweed Symposium: Abstracts, Programs
and Directory. Abstract, Cebu City, Philippines, p.62.
Uyenco, ER., Saniel, L.S., Jacinto, G.S., 1981. The "ice-ice" problem in seaweed
farming. 10th International Seaweed Symposium. Walter de Gruyter, New York,
pp.625-630.
Van der Meer, J.P., 1979. Genetics of Gracilaria sp. (Rhodophyceae,
Gigartinales). V. Isolation and characterization of mutant strains. Phycologia 18, pp.
47-54.
Van Der Meer, J.P., 1990. Genetics. In: Cole, KM., Sheath, R.G. (Eds.), Biology of
the Red Algae. Cambridge Univ. Press, Cambridge, pp.103-122.
Van der Meer, J. P. & X. Zhang, 1988. Similar unstable mutations in three
species of Gracilaria (Rhodophyta). J. Phycol. 24: pp. 198-202.
Velosos, A.R.V., 1989. Cash-Crop from the Sea, SEAFDEC News. 12(2), pp. 5-10.
Wood, W.F., 1989. Photoadaptive responses of the tropical red alga Eucheuma
striatum Schmitz (Gigartinales) to ultra-violet radiation. Aquat. Bot. 33, pp. 41-51.
Vreeland, V., E. Zablackis, R. Doboszewski & W.M. Laetsch, 1987. Molecular
markers for marine algal polysaccharides. Hydrobiologia 151/152, pp. 155-160.
SuriaLink 1-0703

Bibliography Wal-Zer
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.
monograph index
Page 82
next

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Eucheuma Seaplant Production

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SuriaLink Infomedia

Monograph # 1-0703, SuriaLink, July, 2003

The ABC of Eucheuma Seaplant Production

1. Abstract & Index

Photos and diagrams by Iain C. Neish except where noted otherwise

3.Populations 4.A.TaxonomicClassification | 5.B.TaxonomicDescriptions |6.C.Characteristics | 7.D.TradeNames | 8.E.Tambalang |

25.A.PlantElements | 26.B.GeneralPysiology | 27.C.Temperature | 28.D.Light | 29.E.WaterMotion | 30.F.Salinity |

36.A.Overview | 37.B.Background | 38.C.HabitatTypes | 39.D.On-bottom | 40.E.Off-bottom | 41.F.PenCageBagTube |

61.A.ManagingPropagules | 62.B.TLC | 63.C.MovingPropagules | 64.D.BadWeather | 65.E.Productivity | 66.F.Treating |

Abstract: The development of commerce based on eucheuma seaplants is an outstanding

Eucheuma seaplant farm, Bohol, Philippines

The ABC of Eucheuma Seaplant Production

Introduction to the Biology of Eucheuma Seaplants

Farming of the eucheuma seaplants is a continuous process of

Farmers tend to plant and harvest crops on a short cycle of 4-6

The ABC of Eucheuma Seaplant Production

Introduction to the Agronomy of Eucheuma Seaweeds

Success results for a commercial seaplant project only if the best

Note that "success" is defined as being "socially, economically

Crop factors (i.e. agronomic procedures) critical to commercial

Human socio-economic factors germane to the success of commercial

The ABC of Eucheuma Seaplant Production

3.A Taxonomic Classification

This monograph deals with commercially useful genera of the Tribe

The trade name "gelatinae", refers to the scientific species

Phylum Rhodophyta, Class Rhodophyceae, Subclass Florideophycidae, Order

The trade name "spinosum" generally refers to Eucheuma

The trade names, "gelatinae" "spinosum" and "cottonii" and ,

Trade name: gelatinae

Commercial species: gelatinae (GEL)

The specific name "alvarezii" as applied to

Authority: Doty ex P.C. Silva

Type species: Betaphycus philippinensis Doty

Common names: Eucheuma, Gelatinae (also see Eucheuma names)

Type species: Eucheuma denticulatum (N.L. Burman) F.S.Collins & Hervey

Trade name: cottonii

Type species: n/a

The ABC of Eucheuma Seaplant Production

3.B Taxonomic Descriptions

Eucheumatoideae species of commercial significance are notoriously

Species in section Gelatinae tend to grow in very turbulent, active

Many cannot be distinguished on the basis of one specimen or

They are compressed, flattened ultimately with noticeable

They often have long, cylindrical branches (e.g. E. denticulatum) or

The ABC of Eucheuma Seaplant Production

3.C Table of Eucheumatoideae Characteristics

Characteristics of the typical species in the major sections of

1- Fronds of many forms but

1- Fronds cylindrical; spines

1- Fronds compressed; spines

2- Spines in regularly spaced

2- Spines in rows, marginally

2- Spines often scattered, in

3- Branches generally from

3- Branches mostly marginal,

3- Branching generally from

4- Hyphal axial core usually

4- Axial core rhizoidal and

4- Axial core tortuous, hyphal

5- Beta or other carrageenans

6- Cystocarps on main axes

6- Cystocarps on lateral axes

6- Cystocarps on main axes

7- No cystocarp associated with