THEMATIC BACKGROUND STUDY
Genetic Resources for
Farmed Seaweeds
Citationa: FAO. forthcoming. Genetic resources for farmed seaweeds. Rome.
The designations employed and the presentation of material in this information product do
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Contents
List of tables
iii
List of figures
iii
Abbreviations and acronyms
iv
Acknowledgements
v
Abstract
vi
Introduction
1
1.
PRODUCTION, CULTIVATION TECHNIQUES AND UTILIZATION
2
1.1
Species, varieties and strains
2
1.2
Farming systems
7
1.2.1 Sea-based farming
1.2.2 Land-based farming
2.
3.
4.
7
18
1.3
Major seaweed producing countries
19
1.4
Volume and value of farmed seaweeds
20
1.5
Utilization
24
1.6
Impact of climate change
26
1.7
Future prospects
27
GENETIC TECHNOLOGIES
27
2.1
Sporulation (tetraspores and carpospores)
28
2.2
Clonal propagation and strain selection
28
2.3
Somatic embryogenesis
28
2.4
Micropropagation
29
2.4.1 Tissue and callus culture
29
2.4.2 Protoplast isolation and fusion
30
2.5
Hybridization
32
2.6
Genetic transformation
33
MAJOR PROBLEMS OF FARMING SEAWEEDS
36
3.1
Disease and epiphytism
36
3.2
Social and financial
41
IMPACT OF SEAWEED FARMING
41
4.1
Socio-economic impact
41
4.2
Ecological-environmental impact
41
5.
DRIVERS OR MOTIVATIONS TO PURSUE OR EXPAND FARMING
42
5.1
Food
42
5.2
Feed (aquaculture)
42
5.3
Fuel
43
6.
CONSERVATION AND SUSTAINABLE STRATEGIES
43
7.
ENHANCEMENT PROGRAMME
46
7.1
Education
46
7.2
Research and training
46
8.
9.
ROLE OF INTERNATIONAL AND REGIONAL ASSOCIATIONS IN THE DEVELOPMENT
AND MANAGEMENT OF FARMED SEAWEEDS
46
SOURCES OF DATABASES
48
9.1
Regional and international centres
48
9.2
Dissemination, networking and linkages
49
10. EXCHANGE PROGRAMMES
50
10.1 Information
50
10.2 Scientists and experts
51
10.3 Test plants
51
11. CONCLUSIONS
52
References
53
ii
List of tables
1.
Summary of seaweeds currently farmed
3
2.
English and local names of farmed seaweed
6
3.
Summary of the different culture techniques and species farmed by country
8
4.
Organisms suitable for IMTA in temperate waters
15
5.
Selection of sea-based IMTA practices in different countries
16
6.
Selection of land-based IMTA practices in different countries
19
7.
Major seaweed producing countries
20
8.
Major seaweeds farmed in Japan and the Republic of Korea
21
9.
Summary of utilizations of farmed seaweeds
24
10.
Earlier reports on the regeneration of plants from callus
29
11.
Summary of protoplast isolation and regeneration of farmed seaweeds
31
12.
Summary of seaweeds that were hybridize
33
13.
Summary of farmed seaweeds that were genetically transformed
34
14.
Summary of seaweed diseases and epiphytism
37
15.
Conservation and sustainable strategies for farmed seaweeds
45
16.
International, regional and local associations, organizations and societies
engaged in seaweed research and other related activities
47
Some international algae centres
Various networks involved in seaweed farming and allied activities
48
49
17.
18.
List of figures
1
Photos of commercially farmed red seaweeds
4
2
Photos of commercially farmed brown seaweeds
5
3
Photos of commercially farmed green seaweeds
5
4
Examples of sea-based commercial farming
11
5.
Conceptual diagram of an IMTA operation
15
6.
Raceway cultivation of Chondrus crispus at Acadian Seaplants Limited, Canada
18
7.
Ulva grown in tanks/raceways using deep seawater from the Mediterranean
sea (Israel) Ulva fasciata, U. lactuca and U. rigida – Haliotis midae grown in
raceways in Sout
18
Africa.(b) Fish (Sparus aurata) and seaweeds (Ulva and Gracilaria) and
mollusc (Haliotis discus hannai) in Israel at SeaOr Marine Enterpris
18
9.
Seaweed carrageenan (cottonii) production, 2015 (tonnes, dry weight)
21
10.
Seaweed carrageenan (spinosum) production, 2015 (tonnes, dry weight)
22
11.
Gracilaria production by region, 2015 (tonnes, dry weight)
23
12.
Gracilaria production by country, 2014 (tonnes, dry weight)
23
13.
Gelidium production by region, 2015 (tonnes, dry weight)
23
14.
Infection triangle
36
15.
Pyramid schematic of seaweed product markets
42
16.
Sustainability paradigm
42
8.
iii
Acronyms and abbreviations
CO2
carbon dioxide
dwt
dry weight
F1
first generation
F2
second generation
FAO
Food and Agriculture Organization of the United Nations
GUS
glucuronidase
lacZ
bacterial beta-galactosidase
IFREMER
Institut Français de Recherche Pour l’exploitation de la Mer
IMTA
integrated multi-trophic aquaculture
OA
ocean acidification
PGRs
plant growth regulators
Ph.D.
Doctor of Philosophy
PyAct1
P. yezoensis actin1 promoter
PyGUS
P. yezoensis glucuronidase
RM
Malaysian ringgit
SEA
Southeast Asia
SES
Seaweed Energy Solutions
SINTEF
Stiftelsen for Industriell og Teknisk Forskning ved NTH
SV40
a promoter
iv
Acknowledgements
The author is thankful to the following colleagues: Prof. Yusho Aruga of Japan for providing
data on seaweed production; Dr E.K. Hwang of the National Fisheries Research and Development Institute, Republic of Korea, for the latest seaweed production and photos of farmed
seaweeds; and Dr Tong Pang of the Chinese Academy of Sciences Institute of Oceanology and
Prof. Show-Mei Lin for the information provided about the Chinese Phycological Society and the
Taiwanese Phycological Society, respectively.
v
Abstract
The genetic resources of farmed seaweeds are often omitted from the State of the World Report
despite the significance of these seaweeds as a source of human food; a source of natural colloids
as food ingredients for cosmetics, pharmaceutical and nutraceuticals purposes; and a source of
feed in aquaculture. Hence, this review has been done to provide significant data and information on the farmed red, brown and green seaweeds based on the following major areas: (i)
cultivation – species/varieties, techniques, volume and value of production; (ii) genetic technologies; (iii) major problems of farming seaweeds; (iv) drivers or motivations to pursue farming;
(v) conservation and sustainability strategies; (vi) enhancement programmes; (vii) regional and
international collaborations; (viii) sources of databases; and (ix) exchange programmes.
Global seaweed farming occurs predominantly in Asia, both for the brown (Saccharina and
Undaria) and the red seaweeds (Eucheuma, Gelidium, Gracilaria, Kappaphycus and Pyropia),
compared with Europe, which is still small in scale and can be found in countries such as Denmark,
France, Ireland, Norway, Portugal and Spain. Since the beginning, brown seaweeds (Saccharina
and Undaria) dominated the farming of seaweed globally until it was overtaken by red seaweeds in 2010, which come mainly from the Kappaphycus and Eucheuma species. The brown
seaweeds are normally farmed in subtemperate to temperate countries, such as China PR, Japan
and the Republic of Korea, while Kappaphycus and Eucheuma are farmed in subtropical to tropical countries, dominated by Indonesia, the Philippines, Tanzania and Malaysia. At present, 29
species of red seaweeds dominate commercial cultivation, followed by 13 species of brown and
11 species of green seaweeds.
There are other red seaweeds that are currently farmed in open seas or brackish-water ponds, or
in land- based tanks. These are Asparagopsis, Chondrus crispus, Gelidium, Gracilaria, Hydropuntia, Palmaria palmata and Pyropia. Among the green seaweeds, Caulerpa codium, Monostroma
and Ulva are farmed for commercial purposes.
Phyco-mitigation (the treatment of wastes by seaweeds), through the development of integrated
multi-trophic aquaculture (IMTA) systems, has existed for centuries, especially in Asian countries, as a result of trial and error and experimentation. At present, IMTA is a form of balanced
ecosystem management, which prevents potential environmental impacts from fed aquaculture
(finfish) and organic (shellfish) and inorganic (seaweed). It is gradually gaining momentum in
western Europe and Israel. IMTA is mainly aimed towards higher production of seaweed biomass
not only for food and feed purposes, but also as a source of fuel. It also provides exciting new
opportunities for valuable crops of seaweeds.
Traditional selection of strains based on growth performance and resistance to “disease” are still
used in propagating farmed species. The breakthrough in the hybridization of Laminaria japonica in China paved the way to massive cultivation of this species globally. The development of
plantlets from spores for outplanting purposes is still being practised today in some brown (Saccharina and Undaria), red (Palmaria and Pyropia) and green seaweeds (Codium, Monostroma
and Ulva). Micropropagation through tissue and callus culture is becoming a popular method
in generating new and improved strains in Eucheuma and Kappaphycus, though its commercial
use for farming purposes has yet to be tested further. Vegetative propagation is still widely used,
especially in the tropics.
vi
Global climate change is adversely affecting the ecophysiology of farmed seaweeds, leading to
decreased productivity and production, primarily in the tropics. “Diseases” and severe epiphytism are two major technical problems of farmed seaweeds.
The main driver for the continued interest in seaweed cultivation has been the potential for
the production of large volumes of a renewable biomass, which is rich in carbohydrates and
therefore attractive to third-generation biofuel production. Seaweed biomass has a wide range
of applications, such as: (i) biobased and high-value compounds in edible food, food and feed
ingredients, biopolymers, fine and bulk chemicals, agrichemicals, cosmetics, bioactives, pharmaceuticals, nutraceuticals and
botanicals; and (ii) lower-value commodity bioenergy compounds in biofuels, biodiesels, biogases, bioalcohols and biomaterials. Global consumption of sea vegetables is rising as consumers
become more aware of the health and nutritional benefits of these plants. On the other hand,
a biorefinery concept for cultivated seaweed biomass that approaches a complete exploitation
of all the components in the raw material and that creates added value will likely succeed in the
global market in the next few years.
The presence of regional and international networks is of prime importance in the exchange of
databases/information, experts, young scientists and test plants in pursuit of an excellent level
of competency and efficiency in the conduct of projects.
vii
Introduction
The increasing global population needs to source food from the ocean, which is a much greater
area than the land. The ocean is rich with diversified flora and fauna, and both are sources of
protein, vitamins, minerals, phytohormones and bioactive compounds. Thousands of species of
macro-algae (seaweed) dominate the vegetation of the sea floor from the intertidal to the subtidal zone.
The domestication of several economically important seaweed such as Saccharina, Undaria and
Pyropia in China, Japan and the Republic of Korea, and Kappaphycus and Eucheuma in Indonesia,
Malaysia, the Philippines and the United Republic of Tanzania led to intensive commercial cultivation of these seaweeds. Except for the United Republic of Tanzania, the commercial farming of
seaweed, both temperate and tropical species, is centred in Asia. Despite the presence of several
economically important seaweeds in the Western countries, commercial farming is not yet practised there except in a few countries, such as Chile for Gracilaria and Macrocystis (Buschmann et
al., 2001); France for Palmaria palmata, Pyropia umbilicalis and Undaria pinnatifida (Netalgae);
and Canada for Saccharina latissima as integrated multi-trophic aquaculture (IMTA) (Chopin et
al., 2013) and Chondrus crispus, to name a few. Trial farming of Saccharina and P. palmata are
now being cultivated in western Europe, some of which are near the commercial stage.
Seaweeds are farmed mainly for food as sea vegetables and food ingredients (Bixler and Porse,
2011), as well as feed (Wilke et al., 2015; Norambuena et al., 2015). However, Western countries
are seriously looking into biorefinery products from seaweeds, which need a vast amount of
biomass and which must be derived from farming. Sustainability of biomass must come from
farming and not from the harvesting of the natural population.
The world is experiencing climate change, and several reports have shown that seaweeds are an
efficient carbon dioxide (CO2) sink. Seaweed aquaculture beds (SABs) provide ecosystem services
similar to those services gained from seaweed beds in natural or wild habitats. The use of SABs
for potential CO2 mitigation efforts has been established, with commercial seaweed production
in China PR, India, Indonesia, Japan, Malaysia, the Philippines, the Republic of Korea, Thailand
and Viet Nam, and is in the developmental stage in Australia and New Zealand (Chung and Lee,
2014). Seaweed farming is no doubt an aquaculture endeavour that is socially and economically
sustainable (= equitable); socially and environmentally sustainable (= bearable); and economically and environmentally sustainable (= viable) (Circular Ecology, 2016). Every stakeholder has
an important role along the value chain to make it sustainable.
1
1.
PRODUCTION, CULTIVATION TECHNIQUES AND UTILIZATION
For more than 100 years, China PR and other countries in Asia have grown seaweeds (also known
as macro-algae) at a large industrial scale for the production of food, animal feed, pharmaceutical remedies and cosmetic purposes. Commercial cultivation of seaweeds has a long history
in Asia; in fact, the major source of cultivated seaweeds comes from this region. Despite being
described as a low technology, it is highly successful and efficient coupled with intensive labour
at low costs. On the other hand, an emerging rise in investment from petrochemical companies
and governments for projects in Asia, Europe and the Americas aims at extracting sugars from
seaweed for ethanol, bio-based diesel, advanced biofuels, drop-in fuels, biobutanol, biochemical
and biopolymers.
Low technology cultivation practices can become highly advanced and mechanized, requiring on-land cultivation systems for seeding some phases of the life history before grow-out
at open-sea aquaculture sites. Cultivation and seedstock improvement techniques have been
refined over the centuries, mostly in Asia, and can now be highly sophisticated. High technology, on-land cultivation systems have been developed in a few rare cases, mostly in the Western
world, wherein commercial viability can only be reached when high value-added products are
obtained, their markets secured (not necessarily in response to a local demand, but often for
export to Asia), and labour costs reduced to balance the significant technological investments
and operational costs.
1.1
Species, varieties and strains
Among the farmed seaweeds, Chondrus crispus, Eucheuma denticulatum, Kappaphycus alvarezii and K. striatus have different colour morphotypes, which range from brown, green, red,
yellow and purple. Table 1 shows the different genera and species commercially farmed, which
is composed of 11 genera and over 25 species of red seaweeds with two varieties; 7 genera and
12 species of brown seaweeds; and 5 genera and 10 species of green seaweeds with one variety.
Among the red seaweeds, Gracilaria has 11 species, followed by Pyropia with 5 species; in the
brown seaweeds, Sargassum has 4 species; and the green seaweeds are dominated by Ulva with
6 species (Figures 1–3). English and local names of some farmed seaweeds are shown in Table 2.
2
TABLE 1.
Summary of seaweeds currently farmed
Red seaweeds
Genus
Brown seaweeds
Species
Genus
Green seaweeds
Species
Genus
Species
Asparagopsis
armata
Alaria
esculenta
Capsosiphon
fulvescens
Betaphycus
philippinensis
Cladosiphon
okamuranus
Caulerpa
lentillifera
Chondrus
crispus
Hizikia
fusiformis
Eucheuma
denticulatum
Macrocystis
pyrifera
Codium
fragile
Eucheuma
var. milyon milyon
Saccharina
digitata
Monostroma
nitidum
Eucheuma
isiforme
Ulva
compressa
Saccharina
japonica
fasciata
racemosa var. macrophysa
Saccharina
latissima
intestinalis
Gracilaria
asiatica
Sargassum
fulvellum
linza
Gracilaria
changii
Sargassum
horneri
pertusa
Gracilaria
chilensis
Sargassum
muticum
prolifera
Gracilaria
fastigiata
Sargassum
thunbergii
Gracilaria
firma
Undaria
pinnatifida
Gracilaria
fisheri
Gracilaria
heteroclada
Gracilaria
lemaneiformis
Gracilaria
manilaensis
Gracilaria
tenuistipitata
Gracilaria
tenuistipitata var. lui
vermiculophylla
Gelidiella
acerosa
Gelidium
amansii
Hydropuntia
edulis
Kappaphycus
alvarezii
Kappaphycus
malesianus
Kappaphycus
striatus
Palmaria
palmata
Pyropia
dentata
Pyropia
haitanensis
Pyropia
pseudolinearis
Pyropia
seriata
Pyropia
tenera
Pyropia
umbilicalis
3
FIGURE 1.
Photos of commercially farmed red seaweeds. Photos courtesy of EK Hwang, AQ Hurtado
Asparagopsis armata
Chondrus crispus
Palmaria palmata
Kappaphycus alvarezii
Kappaphycus striatus
Eucheuma denticulatum
Eucheuma isiforme
Bataphycus
philippinensis
Gelidium amansii
Gracilaria changii
Graciliaria chorda
Gracilaria firma
Gracilaria heteroclada
Gracilaria tenuistipitata
Gelidiella acerosa
Gracilaria fastigiata
Pyropia dentata
Pyropia haitanensis
Pyropia seriata
4
Pyropia tenera
Pyropia yezoensis
FIGURE 2.
Photos of commercially farmed brown seaweeds. Photos courtesy of EK Hwang, AQ Hurtado
Alaria esculenta
Cladosiphon okamurans
Hizikia fusiformis
Macrocystis sp.
Saccharina digitata
Saccharina japonica
Saccharina latissima
Sargassum fulvellum
Sargassum muticum
Sargassum thunbergii
Undaria pinnatifida
FIGURE 3.
Photos of commercially farmed green seaweeds. Photos courtesy of EK Hwang, AQ Hurtado
Capsosiphon fulvescens
Caulerpa lentillifera
Codium fragile
Monostroma nitidum
Ulva compressa
Ulva fasciata
Ulva intestinalis
Ulva linza
Ulva pertusa
Ulva prolifera
5
TABLE 2.
English and local names of farmed seaweeds
Scientific name
English
Chinese
Japanese
Korean
SEA region
Red
Chondrus crispus
Irish moss
Eucheuma
Spinosum
denticulatum
Gracilaria
Ogonori
Agar-agar
Kappaphycus alvarezii
Tambalang,
besar
Kappaphycus striatus
Elkhorn
Palmaria palmata
Dulse
Pyropia sp.
Purple laver
Flower, sacol
Zicai
Nori
Gim
Hijiki
Tot hiziki
Makombu, Shinori-kombu,
Hababiro-kombu, Okikombu, Uchi kombu, Motokombu, Minmaya-kombu,
Ebisume hirome, Umiyamakombu, Hoiro-kombu,
Kombu
Hae tae,
Tasima
Gamet
Brown
Alaria esculenta
Winged kelp
Hizikia fusiformis
Saccharina digitata
Horsetail kelp
Saccharina japonica
Royal kombu, Japanese
kelp
Saccharina latissima
Sugar kelp, sweet kelp,
sea belt, poor man’s
weather glass, Kombu
royale, sweet wrack,
sugar tang, oarweed
Sargassum muticum
Wireweed
Undaria pinnatifida
Japanese kelp, Asian
kelp, apron-ribbon
vegetable
Hai dai, Hai tai,
Kunpu
Kombu, Kurafuto kombu
Ito-wakame,
Qundai-cai,
Kizami-wakame
Wakame, Ito-wakame,
Kizami-wakami, Nambuwakame
Ito-wakame,
Kizamiwakami,
Miyok
Green
Caulerpa lentillifera
Sea grapes,green caviar
Codium fragile
Green sea fingers, felty
fingers, dead man’s
fingers, stag seaweed,
sponge seaweed, green
sponge, green fleece,
oyster thief, forked felt
alga
Monostroma nitidum
Ulva
Lato
Jiao-mo
Zi-cai
Hitoegusa,
Hirano hitoegusa
Aonori, Aonoriko
Sea lettuce,
green laver
6
1.2
Farming systems
1.2.1 Sea-based farming
Sea-based farming may be classified according to location: (i) coastal; (ii) deep sea; and (iii)
offshore. Coastal and deep-sea farming are common in Asia, and to a little extent in Latin
America and in the western Indian Ocean regions. The fixed off-bottom line, the hanging
longline, single and multiple raft longlines and spider-web techniques of cultivating Eucheuma
and Kappaphycus and sometimes Gracilaria (Figure 4) in the coastal and deep-sea waters are
well documented (Hayashi et al., 2014; Hurtado et al., 2014; Msuya et al., 2014). Gracilaria
and Macrocystis are also commercially farmed in Chile (Buschmann et al., 2001; Gutierrez et
al., 2006). On the other hand, offshore farming is confined to western Europe (Watson, 2014)
and eastern Canada (Chopin and Sawhney, 2009), mainly the monoculture of Saccharina and
Undaria. Seaweed cultivation is currently in its infancy in Europe. Commercial aquaculture
of seaweed is found in France (Brittany, six farms) and Spain (Galicia, two farms), and on an
experimental basis in Ireland, Asturias (Spain), Norway, and the United Kingdom of Great
Britain and Northern Ireland. The main cultivated species are Saccharina latissima and Undaria
pinnatifida. In Ireland, Palmaria palmata farming is being experimented with on the west
coast, but the results seem limited. However, with the fast development of integrated multi-trophic aquaculture (IMTA) as a culture system in Europe, farming of Alaria esculenta, P.
palmata, S. latissima and Laminaria japonica is gaining much attention in this region (Chopin
et al., 2001; Ridler et al., 2007).
China is known as an industry leader in seaweed production and has long experience in
seaweed cultivation, innovation and production. IMTA started in China about 2 000 years ago
with a different system, called spontaneous integrated culture. Most of the culture systems
in the country, however, are still single species intensive culture. China is well known in the
field of marine aquaculture. More than 30 important aquaculture species, including kelp, scallops, oysters, abalone and sea cucumbers, are grown using various culturing methods, such as
longlines, cages, bottom sowing and enhancement, pools in the intertidal zone, and tidal flat
culture (Zhang et al., 2007).
The concept of IMTA was coined in 2004 and refers to the incorporation of species from different
trophic positions or nutritional levels in the same system (Chopin and Robinson, 2004). IMTA,
however, has been successfully practiced in Sanggou Bay in north China since the late 1980s
(Fang et al., 1996). There are several IMTA modes in the bay, with benefits at the ecosystem
level. For instance, the co-culture of abalone and kelp provides combined benefits of a food
source and waste reduction: abalone feed on kelp, and the kelp take up nutrients released from
the abalone (Tang et al., 2013). The co-culture of finfish, bivalves and kelp links organisms from
different trophic levels so that the algae absorb nutrients released from finfish and bivalves and
bivalves feed on suspended fecal particles from the fish. Since kelp and Gracilaria lemaneiformis
are cultured from December to May and from June to November, respectively, nutrients are
absorbed by the algae throughout the year. These examples of multi-trophic culture maximize
the utilization of space by aquaculture as they combine culture techniques in the pelagic and
benthic zones. Implementation of IMTA in Sanggou Bay has improved economic benefits, maintained environmental quality, created new jobs, and led to culture technique innovations (Fang
and Zhang, 2015).
7
Table 3 presents a summary of the different culture techniques of the different farmed seaweeds
per country, all of which are in the commercial stage, with the exception of the land-based IMTA
in Portugal. Apparently, hanging longline is common both to red and brown seaweeds. Except
for Caulerpa, Eucheuma, Gracilaria and Kappaphycus, the source of propagules for commercial
farming comes from spores that are grown first in hatcheries and then outplanted when reaching the juvenile stage during favourable sea temperature. In contrast, these four genera use
vegetative cuttings as propagules for commercial farming.
TABLE 3.
Summary of the different culture techniques and species farmed by country
Country
Red
Brown
Green
Australia
Brazil
Ulva pertusa*1
Gracilaria birdiae*6
Gracilaria domingensis**3
Kappaphycus alvarezii**4,6
Kappaphycus striatus**4,6
Cambodia
Kappaphycus alvarezii**4,6
Kappaphycus striatus**4,6
Canada
Chodrus crispus**1
Alaria esculenta*6
Palmaria palmata*2
Macrocystis integrifolia*6
3
Saccharina latissima*
Caribbean Islands
Gracilaria spp.**6
Chile
Gracilaria chilensis**20,21
Macrocystis pyrifera*6
Betaphycus philippinensis**18
China
Eucheuma denticulatum**4,6
Hizikia fusiformis*6
Gracilaria lemaneiformis**6
Macrocystis pyrifera*10
**13
Gracilaria tenuistipitata var. liui
Saccharina japonica*3
Kappaphycus alvarezii**6
Sargassum fulvellum*6
Kappaphycus striatus**4,6
Sargassum horneri*6
*5
Pyropia haitanensis
Sargassum muticum*6
Pyropia yezoensis*5
Sargassum thunbergii*6
Undaria pinnatifida*3,6
Denmark
Saccharina latissima*2,3
France
Palmaria palmata
Fiji Islands
Kappaphycus alvarezii**6
*1
*2,3
Undaria pinnatifida
Ulva intestinalis*2
Ulva pertusa*2
Kappaphycus striatus**6
India
Eucheuma denticulatum**4,6
Ulva fasciata*5
**5
Gelidiella acerosa
Gracilaria sp.**10
Hydropuntia edulis**1,6
Kappaphycus alvarezii**10
Kappaphycus striatus**10
8
Country
Indonesia
Red
Brown
Green
**4,6
Eucheuma denticulatum
Gracilaria asiatica**13
Gracilaria heteroclada**6,10,13
Gelidium amansii**6
Kappaphycus alvarezii**4,6
Kappaphycus striatus**4,6
Ireland
Asparagopsis armata**6
Alaria esculenta*3
Palmaria palmata*6
Saccharina latissima*3
Israel
Gracilaria sp.
Japan
Gelidium amansii*6
**2
Ulva pertusa**2
Cladosiphon okamuranus*6
*5
*6
Caulerpa lentillifera*8
Pyropia pseudolinearis
Saccharina japonica
Monostroma nitidum*5
Pyropia tenera*5
Undaria pinnatifida*6
Ulva sp.*16
Gracilaria spp.*/**6
Hizikia fusiformis*6
Codium fragile*/**6
Pyropia dentata*5
Saccharina japonica*3
*5
Pyropia yezoensis
Republic of Korea
Capsosiphon
*5
*3
Pyropia seriata
Pyropia tenera*5
*5
Madagascar
Kappaphycus alvarezii**6
Malaysia
Eucheuma denticulatum**6
Kappaphycus alvarezii**6 Kappaphycus
malesianus**6 Kappaphycus striatus**6
Myanmar
Kappaphycus alvarezii**6 Kappaphycus
striatus**6
Saccharina latissima
Ulva compressa*5
Sargassum fulvellum*/**6
Ulva linza*5
*3
Pyropia yezoensis
Undaria pinnatifida
Norway
fulvescens*17
Ulva prolifera*5
Saccharina latissima*3
Panama
Kappaphycus alvarezii**6
Philippines
Eucheuma denticulatum **6
Eucheuma denticulatum var.
milyon milyon**6
Gracilaria changii**10,13
Gracilaria firma**10,13
Gracilaria heteroclada**10,13,14
Gracilaria manilaensis**10,13
Kappaphycus alvarezii**6,7,11,12
Kappaphycus malesianus**6
Kappaphycus striatus**4,6,7,11,12
Caulerpa racemosa var.
Caulerpa lentillifera**14
Caulerpa racemosa var.
macrophysa**
Portugal
Gracilaria vermiculophylla*2
Chondrus crispus*2
Palmaria palmata*2
Codium tomentosum*2
Palmaria palmata*2
Pyropia sp.*2
Caulerpa lentillifera**14
Ulva armoricana*2
Ulva pertusa*2
9
Country
Red
Brown
South Africa
Green
Ulva fasciata**2
Ulva pertusa**2 Ulva
rigida**2
South Pacific
Island
Eucheuma denticulatum**4,6,10
Kappaphycus alvarezii**4,6,10
Solomon Islands
Kappaphycus alvarezii**4
Spain
Palmaria palmata**7
Sri Lanka
Kappaphycus alvarezii**10 Kappaphycus
striatum**10
Tanzania
Eucheuma denticulatum**4 Kappaphycus
alvarezii**10
Taiwan
Gracilaria confervoides**19
Pyropia sp.*5
Caulerpa lentillifera**14
Thailand
Gracilaria fisheri**6,13,14
Gracilaria tenuistipitata**6,13, 14
Hydropuntia edulis**13
Caulerpa lentillifera**2
Undaria pinnatifida*3
Monostroma sp.
Chaetomorpha sp.**19
Ulva sp.**13
Venezuela
Kappaphycus alvarezii**4,6 Kappaphycus
striatus**4,6
Viet Nam
Eucheuma denticulatum**6
Gracilaria asiatica**13,14
Gracilaria firma**13,14 Gracilaria
heteroclada**13,14 Gracilaria
tenuistipitata**13,14 Kappaphycus
alvarezii**6,9 Kappaphycus striatum**6,9
Caulerpa lentillifera**14
United Kingdom
(Scotland)
Alaria esculenta*3
Laminaria digitata*3 Laminaria
hyperborea*3 Saccharina
latissima*3
United States
of America
Pyropia sp.*2
Saccharina latissima*3
Note: *spore; **vegetative.
1
land-based raceways/tanks; 2land-based IMTA; 3sea-based longlines IMTA; 4fixed off-bottom; 5floating nets; 6hanging longline (horizontal); 7hanging
longline (vertical); 8hanging longline (basket bag); 9hanging longline (net bags); 10single raft longline; 11multiple raft longline; 12multiple longline
(spider web); 13pond broadcasting; 14pond “rice-planting”; 15intertidal “rice planting”; 16pole system; 17bamboo-net; 18stone tying; 19co-culture with
shrimps; 20direct burial method; 21plastic tube method.
10
FIGURE 4 (A-T).
Examples of sea-based commercial farming
(a, b) Saccharina japonica on ropes in north west China (photo courtesy of Dr XL Wang)
(c) Saccharina digitata on lines in Ireland
(Watson et al., 2012)
(d) Saccharina latissima on lines in France (photo
courtesy www.c-weed-culture.com)
(e,f) Undaria cultivation in Korea (Kim et al., 2017)
11
(g,h) Hizikia fusiformis cultivation on ropes in Korea (Photos courtesy of EK Hwang )
(i, j) Pyropia net culture (photo courtesy of Yang)
(k) Raft cultivation of Gracilaria in Indonesia
(photo courtesy of S Kusnowirjono)
(i) Palmaria palmata on lines in Ireland (Watson
et al., 2012)
(m) Kappaphycus alvarezii on long lines in
Vietnam (photo courtesy AQ Hurtado)
(n) Kappaphycus alvarezii on rafts in Sri Lanka
(photo courtesy of S Bondada)
12
(o, p) Codium fragile cultivation in Korea using long lines (Hwang et al., 2009)
(q) Capsosiphon net cultivation in Korea
(r) Ulva net cultivation in Korea
(s, t) Monostroma nitidum net cultivation in Japan
13
One of the most discussed types of aquaculture in western Europe, eastern Canada and the
United States of America is IMTA, which is the farming, in proximity, of several species at different trophic levels (Figure 5). The species selected should be well adapted to these conditions
and be appropriately chosen at multiple trophic levels, based on their complementary functions
in the ecosystem as well as for their existing, or potential, economic value. Proximity should be
understood as not necessarily considering absolute distances, but connectivity in terms of ecosystemic functionalities in which management at the sea-area level is paramount.
IMTA is an ecologically engineered ecosystem management approach, which, in fact, does
nothing more than mimic a simplified natural trophic network. IMTA creates a balanced
system for increased environmental sustainability (ecosystem services and green technologies
for improved ecosystem health); economic stability (product diversification, risk reduction and
job creation in coastal communities); and societal acceptability (better management practices,
improved regulatory governance, and appreciation of differentiated and safe products). IMTA
programmes, in different states of development and configuration, are taking place in at least
40 countries (Barrington et al., 2009).
IMTA has gained recognition after 16 years of existence in the West and has slowly been developing in other regions. The most advanced IMTA systems, near commercial or at commercial
scale, can be found in the temperate waters of Canada, Chile, China, Israel and South Africa,
for example (Chopin et al., 2008; Barrington et al., 2009). Table 4 presents the genera selected
based on their established husbandry practices, habitat appropriateness, biomitigation ability
and economic life. Developments of IMTA projects have been started in France, Ireland, Japan,
the Republic of Korea, Mexico, Norway, Portugal, Spain, Thailand, Turkey, the United Kingdom
of Great Britain and Northern Ireland (mostly Scotland), and the United States of America (see
Table 5 for sea-based practices and Table 6 for land-based practices) (Barrington et al., 2009).
IMTA offers many advantages compared with the monoculture system (Barrington et al., 2009),
such as:
(i)
Effluent biomitigation: the mitigation of effluents through the use of biofilters (e.g. seaweeds and invertebrates), which are suited to the ecological niche of the farm.
(ii)
Disease control: prevention or reduction of disease among farmed fish can be provided by
certain seaweeds due to their antibacterial activity against fish pathogenic bacteria (Bansemir et al., 2006), or by shellfish that reduce the virulence of infectious salmon anaemia
virus (Skar and Mortensen, 2007).
(iii)
Increased profits through diversification: increased overall economic value of an operation
from the commercial by-products that are cultivated and sold.
(iv)
Increased profits through obtaining premium prices: potential for differentiation of the
IMTA products through ecolabelling or organic certification programmes.
(v)
Improving local economy: economic growth through employment (both direct and indirect) and product processing and distribution.
14
(vi)
Form of “natural” crop insurance: product diversification may offer financial protection
and decrease economic risks when price fluctuations occur, or if one of the crops is lost to
disease or inclement weather.
FIGURE 5.
Conceptual diagram of an IMTA operation, including the combination of fed aquaculture
(e.g. finfish) with organic extractive aquaculture (e.g. shellfish), taking advantage of the
enrichment in particulate organic matter; and inorganic extractive aquaculture (e.g. seaweeds)
taking advantage of the enrichment in dissolved inorganic nutrients (Chopin et al., 2008).
Note: DIN = dissolved inorganic nutrients; POM = particulate organic matter.
TABLE 4.
Organisms suitable for IMTA in temperate waters
Fish
Crustaceans
Seaweeds
Molluscs
Echinoderms
Polychaetes
Anoplopoma
Homarus
Brown:
Argopecteen
Apostichopus
Arenicola
Dicentrarchus
Penaeus
Alaria, Durvillaea,
Choromytilu
Athyonidium
Glycera
Gadus
Ecklonia, Lessonia,
Crassostrea
Cucumaria
Nereis
Hippoglossus
Laminaria,
Haliotis
Holothuria
Sabella
Melanogrammus
Macrocystis,
Mytilus
Loxechinus
Mugil
Saccharina,
Pecten
Paracentrotus
Oncorhynchus
Sacchoriza,
Placopecten
Parastichopus
Paralichthys
Undaria
Tapes
Psammechinus
15
Fish
Crustaceans
Seaweeds
Pseudopleuronectes
Red:
Salmo
Asparagopsis
Scophthalmus
Molluscs
Echinoderms
Polychaetes
Stichopus
Callophylis
Chondracanthus
Chondrus
Strongylocentrotus
Gigartina
Gracilaria
Gracilariopsis
Palmaria
Sarcothalia
Green:
Ulva
Source: Barrington et al., 2009.
TABLE 5.
Selection of sea-based IMTA practices in different countries
Country
Fish / shrimp
Australia
Canada
Molluscs /
invertebrates
Seaweed
Status
Reference/
company
Thunnus maccoyii
Seriola lalandi
Solieria robusta Ecklonia
radiata
E
Wiltshire et al.,
2015
Salmo salar
Saccharina latissima
Alaria esculenta
Mytilus edulis
Chopin &
Robinson, 2004
CSP P
Ridler et al., 2007
Chlamys farreri Crassostrea
gigas Haliotis discus hannai
Patinopecten yessoensis
Scapharca broughtonii
Apostichopus japonicus
Saccharina japonica
Gracilaria lemaneiformis
C
Fang et al., 1996a
&b; Fang et al.,
2016
Laminaria/Gracilaria
E
Jiang et al., 2009
Gracilaria chilensis
Macrocystis pyrifera
C
Salmo salar
Troell et al.,
1997
Denmark
Oncorhynchus mykiss
Saccharina latissima
C
Marinho et al.,
2015
Denmark
Oncorhynchus mykiss
Chondrus crispus
E
Marinho et al.,
2015
Indonesia
Chanos chanos
Litopenaeus vannamei
E
Putro et al., 2015
Indonesia
Grouper Pomfret fish
Red carp
Abalone Lobster
E
Sukiman et al.,
2014
China
Shrimp, finfish
China
Lateolabrax japonicus
Pseudosciaena crocea
Chile
Ostrea plicatula
Kappaphycus alvarezii
Eucheuma cottonii
16
Country
Fish / shrimp
Molluscs /
invertebrates
Seaweed
Status
Reference/
company
Ireland
Salmo salar
Crassostrea gigas Mytilus
edulis
Laminaria digitata Pyropia
sp.
Asparagopsis armata
E
Kraan, 2010
Japan
Pagrus major
Apostichopus japonicus
Laminaria
Undaria
Ulva
E
Japan
Pagrus major
Ulva
E
Hirata et al.,
1994
Norway
Salmo salar
Mytilus edulis
Laminaria
E
Barrington et al.,
2009
Norway
Salmo salar
Mytilus edulis
Gracilaria
E
Haliotis asinina
Caulerpa lentillifera
Eucheuma denticulatum
Gracilaria heteroclada
E
Dicentrarchus labrax
Scophthalmus maximus
Chondrus crispus
Gracilaria bursa- pastoris
Palmaria palmata
E
Dicentrarchus labrax
Chondrus crispus
Philippines
Portugal
United
Kingdom
Salmo salar
United
Kingdom
Salmo salar
USA
Atlantic cod
Handå, 2012
Largo et al., 2016
Matos et al.,
2006
E
Matos et al.,
2006
E
Stirling & Okumu ,
1995
Palmaria palmata
Laminaria digitata
Laminaria hyperborea
Saccharina latissima
Sacchoriza polyschides
E
SAMS-Loch Duart
Limited/West Minch
Salmon
Pyropia spp.
C
Spain
Scophthalmus maximus
Yokoyama, 2013
Gracilaria bursa- pastoris
Palmaria palmata
Mytilus edulis
Psammechinus miliaris
Paracentrotus lividus
Crassostrea gigas Pecten
maximus Psammechinus
miliaris Paracentrotus lividus
Carmona et al.,
2006
Note: CSPP - Commercial Scale Pilot Project; E - Experimental; C – Commercial
Seaweed is a growing category in Europe, although it is far behind Asia, where marine plants are
part of a longstanding traditional culinary culture.
In France, the largest producer of seaweed is Algolesko, which began harvesting seaweed in
May 2014. Interestingly, two of its partners are oyster growers, which, apart from their obvious
expertise in aquaculture, also demonstrates the complementary nature of seaweed culture with
other types of aquaculture. Future aquaculture production will see more IMTA practices, which
optimizes interaction between species while reducing environmental impact, leading to sustainable production systems that will supply healthy sustainable seafood for future generations. The
potential of seaweed for bioenergy production and a strong interest in developing IMTA have
given a new dimension to seaweed aquaculture.
17
1.2.2 Land-based farming
There are only a few successful commercial land-based tanks/raceways of seaweed farming that
have been reported. These are: (i) Chondrus crispus (three different colour morphotypes) in
Canada as sea vegetables (direct source of human food) grown in raceways (Figure 6); (ii) Ulva
pertusa in Israel grown in raceways using deep seawater from the Mediterranean Sea and used in
diversified food preparations such as pasta, salads, drinks, and abalone feed (SEAKURA) (Figure
7); and (iii) Ulva pertusa in South Africa (Figure 8a) grown in raceways basically as the primary
food of abalone (Bolton et al., 2006; Robertson-Anderson et al., 2008), and SeaOr Marine Enterprise in Israel using fish (Sparus aurata) and seaweeds (Ulva and Gracilaria) and mollusc (Haliotis
discus hannai) (Figure 8b).
FIGURE 6.
Raceway cultivation of Chondrus crispus at
Acadian Seaplants Limited, Canada.
Photo curtesy Acadian Seaplants Limited
(www.acadianseaplants.com)
FIGURE 7:
Ulva grown in tanks/raceways
using deep seawater from
the Mediterranean Sea
(Israel)
FIGURE 8A-B.
(a) Ulva fasciata, U. lactuca and U. rigida – Haliotis midae grown in raceways in South Africa
(photo by R.J. Anderson). (b) Fish (Sparus aurata) and seaweeds (Ulva and Gracilaria) and mollusc
(Haliotis discus hannai) in Israel at SeaOr Marine Enterprise
18
TABLE 6.
Selection of land-based IMTA practices in different countries
Country
Fish/shrimp
Canada
Hippoglossus hippoglossus
Chile
Oncorhynchus kisutch
Oncorhynchus mykiss
France
Molluscs/
invertebrates
Seaweed/micro-algae
Stat
us
Reference/
company
Palmaria palmata
E
Corey et al., 2014
Crassostrea gigas
Gracilaria chilensis
C
Buschmann et al.,
1996
Dicentrarchus labrax
-
Cladophora. Ulva
E
Metaxa et al., 2006
France
-
Crassostrea gigas
Ulva sp.
E
Lefebvre et al.,
2000
Ireland
Oncorhynchus mykiss
-
Pyropia dioica Ulva sp.
Israel
Sparus aurata
Haliotis discus hannai
Gracilaria Ulva
-
SeaOr Marine Farm,
Israel
Portugal
turbot
-
Chondrus crispus Gracilaria
bursa-pastoris Palmaria
palmata,
E
Matos et al., 2006
Republic of
Korea
Sebastes shlegeli
Stichopus japonicus
Sargassum fulvellum
E
Kim et al., 2014
South
Africa
-
Haliotis midae
Gracilaria Ulva
C
Bolton et al.,
2006
Spain
Dicentrarchus labrax
Tapes decussatus
Isochrysis galbana
E/C
Borges et al., 2005
Spain
Scophthalmus maximus
-
Tetraselmis suecica
Phaeodactylum tricornutum
-
USA
Hippoglossus stenolepsis
-
Chondracanthus
exasperatus
C
Söliv International
USA
Anoplopoma fimbria
Haliotis discus hannai
Palmaria mollis
C
Big Island Abalone
Corporation
-
-
Hanniffy & Kraan,
2006;
www.thefishsite.com
Note: CSPP - Commercial Scale Pilot Project; E - Experimental; C – Commercial
1.3
Major seaweed producing countries
Except for Chile, which farms Gracilaria and Macrocystis, and the United Republic of Tanzania,
which cultivates Eucheuma, world farming of seaweed mainly comes from Asia (Table 7).
19
TABLE 7.
Major seaweed producing countries
Species
Major countries
Red
Chondrus crispus
Canada
Eucheuma denticulatum
Indonesia, Philippines, United Republic of Tanzania
Gracilaria spp.
China, Chile, Indonesia, South Africa, Viet Nam
Kappaphycus alvarezii, K. striatus
Indonesia, Malaysia, Philippines, United Republic of Tanzania
Pyropia spp.
China, Japan, Republic of Korea
Brown
Saccharina
China, Japan, Republic of Korea
Hizikia fusiformis
Republic of Korea
Undaria
China, Japan, Republic of Korea
Green
Caulerpa lentillifera
Japan, Philippines, Viet Nam
Codium fragile
Republic of Korea
Monostroma nitidum
Japan
Ulva spp.
Japan, Republic of Korea
1.4
Volume and value of farmed seaweeds
As of 2016, recent production data on Saccharina, Undaria and Pyropia from China were not
available. The author communicated with colleagues in academia and industry, but only Japan
and the Republic of Korea responded to the request. Table 8 shows the volume of farmed seaweeds in Japan and the Republic of Korea.
Because Indonesia and the Philippines are the two main producing countries of Kappaphycus
(cottonii) in the world, it can be gleaned from Figure 9 that Indonesia continues to increase its
production, while the Philippines has decreased its production since 2009. The sudden increase of
production in Indonesia since 2008 is mainly due to the opening of new cultivation areas, considering the presence of thousands of islands in the country. However, the country’s productivity is
only 11 tonnes dry weight (dwt) ha-1year-1. Despite the geographic location of the Philippines,
which is prone to several cyclones every year that destroy farming structures and propagules, the
country’s productivity is 18 tonnes dwt ha-1year-1 (Porse and Rudolph, 2017). Malaysia, though it
is within the Coral Triangle and has vast areas suitable for farming, is still struggling to increase
its production. In 2014 and 2015, 26 076 tonnes and 24 533 tonnes of Kappaphycus, respectively,
were produced (Suhaimi, personal communication).
Production of Kappaphycus in other southeast Asian countries, such as Cambodia, China, India,
Myanmar and Viet Nam, and in Latin America are still small at present and data are not available.
20
TABLE 8.
Major seaweeds farmed in Japan and the Republic of Korea
Genus
Japan (2014)
Republic of Korea (2015)
Volume (tonnes)
Volume (tonnes)
Value (US$1 000)
Red
Gracilaria
Pyropia
316 200
4
8
390 196
319 441
28 157
15 227
442 771
78 409
86
256
321 910
70 104
377
9 964
3 895
997
Brown
Hizikia
Saccharina
32 800
Sargassum
Undaria
43 900
Green
Capsosiphon
Codium
Cladosiphon
15 500
Ulva
6 748
Sources: Korea Ministry of Oceans and Fisheries, 2015; Japan Ministry of Agriculture, Forestry and Fisheries, 2014.
FIGURE 9.
Seaweed carrageenan (cottonii) production, 2015 (tonnes, dry weight)
120,000
100,000
80,000
60,000
40,000
20,000
0
1996
2009
Indonesia
2015
Philippines
Source: Porse and Rudolph, 2017
21
The shallow areas in the coastal zone of the United Republic of Tanzania and Zanzibar allow
favourable cultivation of Eucheuma denticulatum; hence, these locations are major producing
areas. Figure 10 shows the latest production of spinosum (common vernacular name of E. denticulatum) in the three major producing countries.
FIGURE 10.
Seaweed carrageenan (spinosum) production, 2015 (tonnes, dry weight) Source: Porse and Rudolph,
2017.
25,000
Volume (MT, dwt)
20,000
15,000
10,000
5,000
0
1996
2009
Indonesia
2015
Philippines
Tanzania and Zanzibar
Gracilaria and Gelidium are two genus of seaweed suitable for the processing of agar. The
former being more appropriate for food applications while the latter for bacteriological and
biotechnological applications.
Gracilaria is an ubiquitous seaweed, which can be found both in tropic and temperate waters,
while Gelidium is more confined to temperate waters. Hence, one would expect that the sourcing of Gracilaria for agar processing purposes is much easier than Gelidium. The capacity of
Gracilaria to grow in euryhaline areas and to regenerate from fragments are characteristics that
favour intensive cultivation from brackish- water to full seawater areas (Hurtado-Ponce et al.,
1992; Hurtado-Ponce, 1993; Hurtado-Ponce et al., 1997).
Asia-Pacific is the largest producing region of Gracilaria, followed by the Americas (mainly in
Chile), and Africa and Europe (Figure 11). A more detailed graph is presented in Figure 12, which
shows the countries that produce Gracilaria. Just like Kappaphycus production, which is led by
Indonesia, the same is recorded for Gracilaria.
22
FIGURE 11.
Gracilaria production by region, 2015 (tonnes,
dry weight) Source: Porse and Rudolph, 2017.
FIGURE 12.
Gracilaria production by country, 2014 (tonnes,
dry weight) Source: Paravano, 2015.
4 000 2 000
3 000 100
6 000
24 000
10 000
50 000
80 000
26 000
22 000
Asia Pacific
Europe
Americas
Africa
China
Chile
Namibia & South Africa
Others
Vietnam
Japan & Korea
Indonesia
The production of Gelidium is led by Africa with 6 000 tonnes, followed by Asia-Pacific (2 500
tonnes), Europe (1 000 tonnes), and the Americas (600 tonnes) Figure 13.
FIGURE 13.
Gelidium production by region, 2015 (tonnes, dry weight) Source: Porse and Rudolph, 2017.
600
2 500
6 000
1 000
Europe
Asia Pacific
23
Americas
Africa
1.5
Utilization
Farmed seaweeds have been mainly used as sources of direct food in Asia for many centuries;
however, in the past two to three decades, Western countries have started including seaweeds
in their diet for health reasons. Several single species have various applications, as reflected in
Table 9. A total of 59 species are currently farmed and dominated by red seaweed (54.3 percent),
followed by brown (23.7 percent), and finally green (22.0 percent). Seaweeds are prime candidates for the integrated biorefinery approach – on the one hand, there is a wide range of biobased, high-value compounds (such as edible food, food and feed ingredients, biopolymers, fine
and bulk chemicals, agrichemicals, cosmetics, bioactives, pharmaceuticals,nutraceuticals, botanicals), and on the other hand, lower-value commodity bioenergy compounds (including biofuels,
biodiesels, biogases, bioalcohols, biomaterials).
TABLE 9.
Summary of utilizations of farmed seaweeds
Species
Food
Feed
Food ingredient
Sea vegetable
Agar
Carrageenan
Fuel*
Alginate
Red
Asparagopsis armata
x
Betaphycus philippinensis
x
Chondrus crispus
x
x
Eucheuma denticulatum
x
x
x
Eucheuma denticulatum
var. milyon milyon
x
x
x
Gelidiella acerosa
x
x
Gelidium amansii
x
x
Gracilaria asiatica
x
x
Gracilaria birdiae
x
x
Gracilaria changii
x
x
x
Gracilaria chilensis
x
x
x
Gracilaria domingensis
x
x
Gracilaria firma
x
x
x
Gracilaria fisheri
x
x
x
Gracilaria heteroclada
x
x
x
Gracilaria lemaneiformis
x
x
Gracilaria manilaensis
x
x
x
Gracilaria tenuistipitata
x
x
x
Gracilaria tenuistipitata
var. liui
x
x
x
x
x
Gracilaria vermiculophylla
Gracilaria sp.
x
x
Hydropuntia edulis
x
Kappaphycus alvarezii
x
x
x
24
x
Species
Food
Feed
Food ingredient
Sea vegetable
Agar
Carrageenan
Kappaphycus malesianus
x
x
Kappaphycus striatus
x
x
Palmaria palmata
x
Pyropia dentata
x
Pyropia haitanensis
x
Pyropia pseudolinearis
x
Pyropia seriata
x
Pyropia tenera
x
Pyropia umbilicalis
x
Pyropia yezoensis
x
Pyropia sp.
x
Fuel*
Alginate
x
x
x
Brown
Alaria esculenta
x
Clad siphon okamuranus
x
Hizikia fusiformis
x
Macrocystis integrifolia
Macrocystis pyrifera
x
x
x
x
Saccharina digitata
x
x
Saccharina hyperborea
x
x
Saccharina japonica
x
x
Saccharina latissima
x
x
Sargassum fulvellum
x
x
Sargassum horneri
x
x
Sargassum muticum
x
x
Sargassum thunbergii
x
x
Undaria pinnatifida
x
x
Green
Capsosiphon fulvescens
x
Caulerpa lentillifera
x
Caulerpa racemosa var.
macrophysa
x
Codium fragile
x
Codium tomentosum
x
Monostroma nitidum
x
Ulva compressa
x
x
Ulva fasciata
x
x
Ulva intestinalis
x
x
Ulva linza
x
x
Ulva pertusa
x
x
Ulva prolifera
x
x
Ulva sp.
x
x
*Experimental stage.
25
x
x
1.6
Impact of climate change
Seaweeds are a key source of carbon in the reef ecosystem, and they are involved in other important processes, including the construction of reef frameworks, coral settlements and creation of
habitats. They are a direct food source for herbivorous fish, crabs and sea urchins. The carbon
they produce in photosynthesis enters the food chain via the microbes.
Seaweeds are subject to both regional and global environmental changes in coastal waters,
where environmental factors fluctuate dramatically because of high biological production and
land runoff. Because global ocean changes can influence coastal environments, global warming-induced ocean warming and ocean acidification (OA) caused by atmospheric CO2 rise and
increasing ultraviolet B irradiance at the earth’s surface thus affect the physiology, life cycles and
community structures of seaweeds. According to Ji et al., (2016), some species tested showed
enhanced growth and/or photosynthesis under elevated CO2 levels or ocean acidification conditions, possibly due to increased availability of CO2 in seawater with neglected influence of pH
drop. Nevertheless, OA can harm some macro-algae because of their high sensitivity to the acidic
perturbation to intracellular acid-base stability. Mild ocean warming has been shown to benefit
most macro-algae examined. OA may positively affect gametogenesis because of increased
availability of CO2 and may neutrally influence germination due to the counteractive effects of
decreased pH (Roleda et al., 2012). OA can impact photosynthesis and respiration differently in
some macro-algae. While it is important to look into responses of macro-algae to diel fluctuating
pH (common in coastal waters) under OA (Cornwall et al., 2012), the impacts of OA would affect
productivity of sea-farmed macro-algae that experience dramatic diel pH variations.
Increased availability of carbon and increased acidity in seawater with atmospheric CO2 rise
may have counteractive effects on the physiological activities and growth of macro-algae, and
altered chemistry under OA may reduce growth, photosynthesis and even lead to death of some
macroalgal species (Israel and
Hophy, 2002; Martin and Gattuso, 2009). Ultraviolet B, which penetrates only several metres in
coastal waters, is harmful for macro-algae either in their adult stages or throughout their life
cycles.
Sea level rise may create more available habitat space for macro-algae to grow, as more land
area will be inundated with water. While the increase could impact some species that live in
shallow habitats by reducing their exposure to sunlight (the more water will mean more distance
for sunlight to travel to reach the macro-algae), as a group macro-algae is not vulnerable to negative impacts of sea level rise. The predicted increase in the frequency of severe weather events
such as cyclones, storms and floods will bring an influx of nutrients into the reef ecosystem,
which will increase macro-algae growth and reproduction. Cyclones and storms can also destroy
coral reef structures, increasing habitat areas for macro-algae to grow.
Rising sea temperatures will increase the production of some species of macro-algae. Changes
in temperatures could also lead to changes in these species’ life cycles, growth and production.
Climate change is more notably felt in the tropics than in the temperate countries. The most
notable impact of rising temperature and a concomitant elevated salinity have been reported
on farmed Kappaphycus. The high incidence of “ice-ice” (a disease affecting Kappaphycus and
26
Eucheuma production) as well as epiphytic filamentous algae, were reported in southeast Asia
by Critchley et al. (2004), Hurtado and Critchley (2006), Vairappan (2006), Vairappan et al., (2008),
Tisera and Naguit (2009), Borlongan et al. (2011); in China by Pang et al., (2011, 2012, 2015); and
in Madagascar by Ateweberhan et al., (2015) and Tsiresy et al., (2016).
Low productivity and production and the unavailability of propagules for the next growing
cycles were the major problems of seaweed farmers as a result of rising temperatures. Sometimes the seaweed farmers stopped cultivating Kappaphycus, and consequently, their economic
life was severely affected. While there are possible positive effects of ocean warming for some
warm seawater-grown species, for the cold seawater-grown species, the rise of temperature may
reduce their living space and narrow their available niche.
1.7
Future prospects
Farmed seaweeds in the tropics and subtropics will continue to grow and expand, not only
because of their economic significance among coastal fishers, but also because of the opening
and discovery of more product applications in food industries as well as in pharmaceuticals,
nutraceuticals, cosmetics and personal care. The combination of increasing production, innovative products and consumer demand for natural and organic products will no doubt lead to
bright days for seaweed in Europe and other parts of the globe.
In the temperate countries, especially in western Europe, northeastern Canada and the United
States of America, the brown seaweed Alaria, Laminaria and Saccharina will be expanded tremendously in terms of sea cultivation, both as a monocrop and as part of the IMTA mainly for
biorefineries. Further, sea vegetables like Chondrus crispus, Palmaria palmata, Pyropia yezoensis
and Ulva pertusa will be cultivated extensively in land-based systems both as a monoculture and
IMTA.
IMTA will find its way in countries where intensive fish cage and pond shrimp farming are practised, as in southeast Asia, India and South America. IMTA is considered more sustainable than
the common monoculture systems – a system of aquaculture where only one species is cultured
– in that fed monocultures tend to have an impact on their local environments due to their
dependence of supplementation with an exogenous source of food and energy without mitigation (Chopin et al., 2001). For some twenty years now, many authors have shown that this
exogenous source of energy (e.g. fish feed) can have a substantial impact on organic matter and
nutrient loading in marine coastal areas (Gowen and Bradbury, 1987; Folke and Kautsky, 1989;
Chopin et al., 1999; Cromey et al., 2002), affecting the sediments beneath the culture sites and
producing variations in the nutrient composition of the water column (Chopin et al., 2001).
2.
GENETIC TECHNOLOGIES
The global seaweed industry produced 23–24 million tonnes of wet seaweed from aquaculture
in 2012 (FAO, 2014), as the demand for seaweed based-products exceeds the supply of seaweed
raw material from natural stocks. Aquaculture of seaweed offers advantages over the harvest
of natural stocks for the following reasons: (i) stable supply and reliable access of raw material;
(ii) uniformity of quality of the raw material; and (iii) facilitates the selection of germplasm with
27
desired traits. Seaweed cultivation must be technically feasible, environmentally friendly, economically equitable, and socially acceptable in order to be sustainable.
Traditional selection of strains based on growth performance and resistance to “disease” are still
used in propagating farmed species. The breakthrough in the hybridization of Laminaria japonica in China paved the way to massive cultivation of this species globally. In vitro cell culture techniques have also been employed, as these facilitate development and propagation of genotypes
of commercial importance. There are more than 85 species of seaweeds for which tissue culture
aspects have been reported.
Initially, the aim of these techniques focused mostly on genetic improvement and clonal propagation of seaweeds for mariculture; however, recently, the scope of these techniques has been
extended for use in bioprocess technology for the production of high-value chemicals of great
importance in pharmaceuticals and nutraceuticals, and more recently, in biorefinery.
2.1
Sporulation (tetraspores and carpospores)
All brown seaweeds commercially cultivated (Hizikia, Macrocystis, Saccharina and Undaria) use
strings for the attachment of zoospores in hatcheries during summertime until they reach 1 mm
long, and then they are outplanted into the sea in autumn. When these individuals attain a size
of more than 1 m long, they are ready to harvest. The growth stage from the land-based hatchery to grow-out is nine to ten months.
A number of reports have been conducted on the trial use of spores from Gracilaria for possible
commercial cultivation, but as of 2016 no one has adopted the use of spores for commercial
propagation. Likewise, the use of carposporelings from Kappaphycus alvarezii as possible propagules for field cultivation (Azanza and Aliaza, 1999; Azanza-Corrales et al., 1996; Azanza and
Ask, 2003) did not gain much success compared with the carposporelings from K. striatus, which
were field cultivated in Guimaras Island, the Philippines (Luhan and Sollesta, 2010). Further,
the use of tetrasporelings from K. alvarezii (de Paula, 1999; Bulboa et al., 2007) also did not
gain much attention among the seaweed farmers to use in commercial cultivation compared
with other species, such as Laminaria digitata, Palmaria palmata, Pyropia yezoensis, Saccharina
latissima and Undaria pinnatifida. This is probably because of the low germination rate under
laboratory/hatchery conditions for mass field cultivation. Hatchery production of the conchocelis
and/or spores for outplanting purposes is already well developed in China, Japan and the Republic of Korea and is still practised today.
2.2
Clonal propagation and strain selection
Clonal propagation is the most common and simplest approach to select superior strains from wild
populations to improve the performance of cultivated crops (Santelices, 1992), such as Chondrus
(Cheney et al., 1981), Gigartina (Sylvester and Waaland, 1983), Gracilaria (Patwary and van der Meer,
1982, 1983), and Kappaphycus (Doty and Alvarez, 1973). These studies exploited the organogenetic
potential of seaweeds in isolating superior clones for cultivation. Clonal propagation of Chondrus
crispus in raceways in Canada is the only known successful cultivation of this red seaweed. Its commercial cultivation has been perfected after more than ten years of trial cultivation.
2.3
Somatic embryogenesis
Somatic embryogenesis is an asexual form of plant propagation that mimics many of the events
of sexual reproduction. Also, this process may be reproduced artificially by the manipulation of
28
tissues and cells in vitro. Some of the most important factors for a successful plant regeneration
are the culture medium and the environmental incubation conditions. In vitro somatic embryogenesis is an important prerequisite for the use of many biotechnological tools for genetic
improvement as well as for mass propagation.
Whole plants are regenerated from culture via two different processes: (i) somatic embryogenesis, in which cells and tissues develop into a bipolar structure containing both root and shoot
axes with a closed vascular system (essentially, the type of embryogenesis that occurs in a seed);
and (ii) organogenesis, in which cells and tissues develop into a unipolar structure, namely a
shoot or a root with the vascular system of this structure often connected to parent tissues.
2.4
Micropropagation
2.4.1 Tissue and callus culture
Tissue culture is the science of maintaining cells and/or tissues in vitro in a sterile environment
that regulates specific growth and development patterns. Culture conditions requiring
control include: (i) physical conditions (controlled with an environmental chamber or walk-in
culture room), light, temperature, photoperiod and aeration; and (ii) chemical conditions (controlled by the culture media) – all essential nutrients, minerals, pH and quality of water. Culture
media is either solid (agar) or liquid. Plant growth regulators (PGRs) are essential to induce
developmental changes in cells to create specific tissues. There are five classes of PGR, namely:
(i) auxins – promote both cell division and cell growth; (ii) cytokinins – promote cell division; (iii)
gibberellins – for cell division; (iv) abscisic acid – inhibits cell division; and (v) ethylene – controls
fruit ripening.
Plants can be regenerated in tissue culture either from tissue explants or from isolated cells.
When plant cells and tissues are cultured in vitro, in most cases they exhibit a very wide range
of plasticity. Regeneration of the whole plant from any single cell depends on the concept that
each cell, if given the appropriate stimuli, has the genetic potential to divide and differentiate
into all types of tissues. This genetic potential by plant cells is referred to as totipotency. Several
species of red, brown and green macro-algae have been reported to regenerate from callus, as
shown in Table 10. Although several successful studies were reported on the regeneration of
plantlets of Kappaphycus and Euchuema from callus through micropropagation using different
culture media, their economic viability in the field has yet to be tested further, though initial
trials have been started.
TABLE 10.
Earlier reports on the regeneration of plants from callus
Species
Status of success
Major media and PGR used
Reference
Red
Chondrus crispus
Plant development
SWM3
Chen & Taylor, 1978
Eucheuma sp.
Callus formation
PES
Polne-Fuller & Gibor, 1987
E. denticulatum
Plant development
ESS + IBA and kinetin
Dawes & Koch, 1991
Plant development
ESS + IBA and kinetin
Dawes et al., 1993
Plant development
ESS/2 + PAA and kinetin
Hurtado & Cheney, 2003
Gelidium sp.
Plant development
SSW + NH4NO3 + (NH4)2HPO4
Titlyanov et al., 2006a
Gracilaria changii
Plant development
mES; PES
Yeong et al., 2008
G. tenuistipitata
Palnt development
PGRs
Yokoya et al., 2004
29
Species
Kappaphycus alvarezii
Status of success
Major media and PGR used
Reference
Plant development
ESS + IBA and kinetin
Dawes & Koch, 1991
Plant development
ESS + IBA and kinetin
Dawes et al., 1993
Plant development
PES + NAA, BA, spermine
Munoz et al., 2006
Plant development
ESS/2 + PAA and kinetin
Hurtado & Biter, 2007
Plant development
AMPEP + PAA and kinetin
Hurtado et al., 2009; Yunque et
al., 2011
Plant development
VS 50, f/2 50, ASP12-NTA + IAA,
2-4-D, BA and
colchicine
Hayashi et al., 2008
Plant development
PES, VS 50, F/2 + IAA and BAP
Yong et al., 2014
Plant development
VS 50 + IAA, kinetin, spermine,
colchicine or oryzalin
Neves et al., 2015
Callus formation
VS 50, f/2 50, ASP12-NTA
Zitta et al., 2013
Callus formation
PES + IBA + 6-BA
Li, et al., 2015
Callus and filament
formation
PES and Conway + BA + IAA; BA +
NAA
Sulistiani et al., 2012
Plant development
PES + BAP, NAA, NSE
Yong et al., 2014
Plant regeneration
KTH f/2
Titlyanov et al., 2006b;
Sanderson, 2015
Laminaria japonica
Plant regeneration
MS + Vit. B2 + C-751
Yan, 1984
Undaria pinnatifida
Plant regeneration
MS + Vit. B2 + C-751
Zhang, 1982; Yan, 1984;
Kawashima & Tokuda, 1993
Callus induction
PES
Polne-Fuller & Gibor, 1987
Palmaria palmata
Brown
Green
Ulva intestinalis
2.4.2 Protoplast isolation and fusion
Protoplasts are living plant cells without cell walls that offer a unique uniform single cell system
that facilitates several aspects of modern biotechnology, including genetic transformation
and metabolic engineering. Protoplasts isolation from macrophytic benthic marine algae was
reported as early as 1970 using mechanical methods (Tatewaki and Nagata, 1970; Enomoto and
Hirose, 1972; Kobayashi, 1975). However, the success in producing a large number of viable
protoplasts became possible only after the development of an enzymatic method by Millner et
al. (1979) for Enteromorpha intestinalis (Linnaeus) Nees. Plantlet regeneration from the same
species was reported by Rusing and Cosson (2001).
Only a few species among the farmed seaweeds were tested for protoplast isolation and its
possible regeneration to plantlets. Among the brown seaweeds, only Laminaria japonica (Saga
and Sakai, 1984, Tokuda and Kawashima, 1988; Sawabe et al., 1993; Sawabe and Ezura,
1996; Inoue et al., 2008); L. saccharina and L. digitata (Butler et al.,1989); Macrocystis pyrifera
(Kloareg et al., 1989); and Undaria pinatifida (Tokuda and Kawashima, 1988) were reported.
Only the works of Kloareg et al., (1989) on M. pyrifera and Matsumura et al., (2000) on L. japonica were successful in the regeneration of plantlets from protoplasts.
30
Early protoplast isolations from Kappaphycus alvarezii were made with the purpose of improving the genetic characteristics of this species as a source of propagules for possible commercial cultivation (Zablackis, et al., 1993). Digestions with cellulase and kappa-carrageenase produced only a few cortical cell protoplasts, while digestions with cellulase and iota-carrageenase
only produced epidermal cell protoplasts. When both carrageenases were used in the digestion
media with cellulase, protoplasts were released from all cell types and yields ranged from 1.0 to
1.2 × 107 cells g-1 with sizes from 5 to 200 mm diameter. Protoplasts were subsequently cultured
to study cell wall regeneration; however, no regeneration of plantlets was observed.
Attempts to isolate protoplast from tissue fragments (<1 mm2) of three Philippine cultivars of
Kappaphycus alvarezii, namely the giant cultivar, the cultivar L and the Bohol wild type, by
enzymatic dissolution of cell walls was reported by Salvador and Serrano (2005). The yields of
viable protoplasts from young and old thalli (apical, middle, basal segments) were compared
at various temperatures, duration of treatment and pH using eight combinations of commercial enzymes (abalone acetone powder and cellulase), and prepared extracts from fresh viscera
of abalone (Haliotis asinina) and a terrestrial garden snail. Though viable protoplasts formed
radially expanded discs and filaments arising from the disc, no regeneration to a plantlet was
reported. Table 11 shows a summary of earlier reports on protoplast isolation and regeneration.
As of 2016, protoplast isolation and regeneration is not being used commercially and all applications remain in the research and development phase.
TABLE 11.
Summary of protoplast isolation and regeneration of farmed seaweeds
Species
Status
Reference
Red
Gelidium robustum
PI
Coury et al., 1993
Gracilaria asiatica
PI
Yan & Wang, 1993
Gracilaria changii
PI
Yeong et al., 2008
G. chilensis
PR
Cheney, 1990
G. gracilis
PI
Huddy et al., 2013
G. tenuistipitata
PI
Chou & Lu, 1989; Bjork et al., 1990
Kappaphycus alvarezii
PI
Zablackis et al., 1993; Salvador & Serrano, 2005
Palmaria palmata
PI
Liu et al., 1992; Nikolaeva et al., 1999
Pyropia tenera
PI
Song & Chung, 1988; Fujita & Saito, 1990
P. yezoensis
PI
Fujita & Saito, 1990
P. yezoensis
PR
Yamazaki, et al., 1998; Hafting, 1999
Cladosiphon okamurans
PR
Uchida & Arima, 1992
Laminaria digitata
CW
Butler et al., 1989
L. digita
PR
Benet et al., 1997
L. japonica
PI
Saga & Sakai 1984; Sawabe & Ezura, 1996; Sawabe et al., 1997; Matsumura et al.,
2000
L. saccharina
CW
Butler & Evans, 1990
L. saccharina
PI
Benet et al., 1994
L. saccharina
PR
Benet et al., 1997
Brown
31
Species
Status
Reference
Monostroma nitidum
PI
Yamaguchi et al., 1989
Monostroma nitidum
PR
Fujita & Migita, 1985; Uppalapati & Fujita, 2002
Ulva fasciata
PR
Chen & Shih, 2000
U. flexuosa
PR
Reddy et al., 2006
U. intestinalis
PR
Rusing & Cosson, 2001; Millner et al. 1979
U. pertusa
PI
Saga, 1984; Yamaguchi et al., 1989
U. pertusa (wild)
PI
Reddy et al., 2006; Yamaguchi et al., 1989
Green
U. pertusa (wild)
PR
Chou & Lu, 1989; Reddy et al., 2006
U. pertusa (mutant)
PR
Zhang, 1983; Fujimura et al., 1989; Reddy et al., 1989; Uchida et al., 1992;
Uppalapati & Fujita, 2002
Note: CW = cell wall formation; PI = protoplast isolation; PR = plant regeneration.
2.5
Hybridization
Among the commercial farmed seaweeds, only a few brown and red species have been subjected
to hybridization attempts. The first seaweed that was successfully hybridized was Saccharina (=
Laminaria) done by Chinese scientists during the late 1950s and early 1960s. China pioneered
the method of hybridization, considering that Saccharina japonica was an introduced species
from Hokkaido, Japan. This hybrid Saccharina created a few highly productive strains that were
partially responsible for the increasing annual production in China.
One successful farming of seaweed recorded as a result of hybridization is S. japonica in China.
This seaweed was bred by crossing gametophytes and self-crossing the best individuals and
selecting the best self-crossing line (Li et al., 2016). Its sporophytes were reconstructed each year
from representative gametophyte clones, from which seedlings were raised for farming. Such
strategy ensured Dongfang No. 7 against a variety of contamination due to cross-fertilization,
and occasional mixing and inbred depletion due to self-crossing number-limited sporophytes
matured year after year. Dongfang No. 7 is derived from an intraspecific hybrid through four
rounds of self-crossing and selection and retains a certain degree of genetic heterozygosity, and
thus is immune to inbred depletion because of diversity reduction. Most importantly, farming
Dongfang No. 7 was compatible when used in the farming system. It increased the air dry yield
by 43.2 percent over two widely farmed controls on average, close to the increased intraspecific
hybrid, but less than that of interspecific hybrids or the varieties derived from them. Such strategy was feasible at least for genetically improving the brown algae with a similar life cycle, e.g.
Undaria pinnatifida and Macrocystis pyrifera.
The successful work of Hwang et al. (2014) on the hybridization of female U. pinnatifida and
male U. peterseniana led to the extended period of availability of Undaria for abalone feed
and cultivation in the Republic of Korea. Using free-living gametophyte seeding and standard
on-growing techniques, the second generation (F2) hybrids were found to have longer pinnate
blades and narrower midribs than the first generation (F1) hybrid and formed only sporophylls.
The growth and morphology of F2 hybrids originating from the sporophyll or sorus of the F1
hybrids were not morphologically different from each other. Both of the F2 hybrids exhibited late
maturation, with the early stages of sporophylls appearing in April.
32
An attempt to hybridize Kappaphycus alvarezii and Eucheuma denticulatum was successful, as
reported by Wang (1993), using a somatic cell-fusion method to produce hybrids of non-filamentous or anatomically complex algae as evidenced by isoenzyme electrophoresis. However, this
was not pursued further for its mass production for possible commercial cultivation (Table 12).
TABLE 12.
Summary of seaweeds that were hybridized
Fusion species
Status
Reference
Red
Gracilaria chilensis × G. tivahiae
Plant development
Cheney, 1990
Porphyra yezoensis (red) × P. yezoensis (green)
Plant development
Fujita & Migita, 1987
P. yezoensis × P. pseudolinearis
Plant development
Fujita & Saito, 1990
P. yezoensis × P. haitanensis
Callus development
Dai et al., 1993
P. yezoensis × P. tenera (green)
Callus development
Araki & Morishita, 1990
P. yezoensis (green) × P. suborbiculata
Callus development
Mizukami et al., 1995
P. yezoensis × P. vietnamensis
Callus development
Matsumoto et al., 1995
P. tenera × P. suborbiculata
Callus development
Matsumoto et al., 1995
P. yezoensis × Bangia atropurpurea
Callus development
Fujita, 1993
P. yezoensis × Monostroma nitidum
Plant development
Kito et al., 1998
Ulva pertusa × U. conglobata
Plant development
Reddy & Fujita, 1989
U. pertusa × U. prolifera
Plant development
Reddy et al.,1992
Ulva × Pyropia yezoensis
Protoplast fusion
Saga, et al., 1986
U. linza × U. Pertusa
Protoplast fusion
Jie, 1987
Sporeling production
Shan et al., 2013
Green
Brown
Undaria pinnatifida
(female gametophyte, from parthenosporophytes,
× male gametophyte)
2.6
Genetic transformation
Genetic transformation occurs at the cellular level and can be used to introduce trait altering
genes into the host genome. Cells must be regenerated into plants to recover the transgenic
plant. Genetic transformation is a powerful tool not only for elucidating the functions and regulatory mechanisms of genes involved in various physiological events, but also for establishing organisms that efficiently produce biofuels and medically functional materials, or that carry
stress tolerance under uncertain environmental conditions (Torney et al., 2007; Bhatnagar-Mathur et al., 2008). As of 2016 no genetically transformed seaweeds are being sold or used commercially for food, biofuel or any other applications; this technology is only used for research
and development purposes.
Donald P. Cheney is the pioneer in researching red algal transformation. He and his colleague
performed transient transformation of the red alga Kappaphycus alvarezii using particle bombardment, which was the first report about the transient transformation of seaweeds (Kurtzman
and Cheney, 1991). Since then, there have been recent developments in macroalgal transforma-
33
tion. The report of Wang et al., (2010a) showed a viable way of producing stable transformants
to eliminate chimeric expression, and to achieve transgenic breeding in K. alvarezii using SV40
promoter-driving lacZ gene into cells of K. alvarezii through particle bombardment of epidermal
and medullary cells at 650 psi (pounds per square inch) at a distance of 6 cm. In another report,
a transgenic K. alvarezii was successfully produced when a binary vector pMSH1-Lys carrying
a chicken lysozyme (Lys) gene was transformed into Agrobacterium tumefaciens LBA4404 by
triparental mating (Handayani et al., 2014). The percentage of pMSH1-Lys transformation on K.
alvarezii was 23.5 percent, while the efficiency of regeneration was 11.3 percent. PCR analysis
showed that three of the regenerated thallus contained the lysozyme gene, which has the ability
to break down the bacterial cell wall, a significant result in the prevention of “ice-ice” disease
in K. alvarezii.
Among the red industrially important macro-algae such as Chondrus, Gelidium, Kappaphycus
and Pyropia, the transient gene expression system has not yet been developed in these red macro-algae other than P. yezoensis. Optimization of codon usage in coding regions of the reporter
gene and recruitment of endogenous strong promoters (pPyAct1-PyGUS and pPyAct1-GUS plasmids) are important factors in the transient gene expression system. Furthermore, the use of
particle bombardment is the proven method of gene transfer into red algal cells (Mikami et al.,
2011) (Table 13).
TABLE 13.
Summary of farmed seaweeds that were genetically transformed
Species
Status of
expression
Method of gene
transfer
Promoter
Marker or
reporter
Reference
Gracilaria changii
Stable
Particle bombardment
SV40
lacZ
Gan et al., 2004
Gracilaria changii
Transient
Particle bombardment
SV40
lacZ
Gan et al., 2003
Kappaphycus alvarezii
Transient
Biolistic particle
CaMV 35S
GUS
Kurtzman & Cheney,
1991
Kappaphycus alvarezii
Stable
Particle bombardment
SV40
lacZ
Wang et al., 2010a
Pyropia haitanensis
Stable
Glass bead agitation
SV40
lacZ; EGFP
Wang et al., 2010b
P. tenera
Transient
Electroporation
CaMV 35S
GUS
Okauchi & Mizukami,
1999
P. tenera
Transient
Particle bombardment
PtHSP70; PyGAPDH
PyGUS
Son et al., 2012
P. yezoensis
Transient
Electroporation; particle
bombardment
CaMV 35S
GUS
Kuang et al., 1998
Red
P. yezoensis
Transient
Electroporation
rbcS
GUS
Hado et al., 2003
P. yezoensis
Transient
Electroporation
CaMV 35S
GUS
Liu et al., 2003
P. yezoensis
Transient
Electroporation
CaMV 35S;
B-tubulin
GUS
Gong et al., 2005
P. yezoensis
Transient
Electroporation
CaMV 35S
CAT, GUS
He et al., 2001
P. yezoensis
Transient
Electroporation
Rubusico
GUS, sGFP;
(S65T)
Mizukami et al., 2004
P. yezoensis
Transient
Particle bombardment
CaMV 35S;
PyGAPDH
PyGUS
Hado et al., 2003
P. yezoensis
Transient
Particle bombardment
PyAct1
PyGUS
Takahashi et al., 2010
P. yezoensis
Transient
Particle bombardment
PyAct1
AmCFP; ZsGFP
Mikami et al., 2009
34
Species
Status of
expression
Method of gene
transfer
Promoter
Marker or
reporter
Reference
P. yezoensis
Transient
Particle bombardment
PyAct1
ZsYFP, sGFP
(S65T)
Uji et al., 2010
P. yezoensis
Transient
Particle bombardment
PtHSP70; PyGAPDH
PyGUS
Son et al., 2012
P. yezoensis
Stable
Agrobacterium- mediated
gene transfer
Unknown
Unknown
Bernasconi et al.,
2004
P. yezoensis
Stable
Agrobacterium- mediated
gene transfer
CaMV 35S
GUS
Cheney et al., 2001
Laminaria japonica
Transient
Particle bombardment
CaMV 35S
GUS
Qin et al., 1998
L. japonica
Stable
Particle bombardment
SV40
GUS
Jiang et al., 2003
L. japonica
Transient
Particle bombardment
CaMV 35S, UBI,
AMT
GUS
Li et al., 2009
Brown
L. japonica
Stable
Particle bombardment
FCP
GUS
Li et al., 2009
L. japonica
Stable
Particle bombardment
SV40
HBsAg
Jiang et al., 2002
L. japonica
Stable
Particle bombardment
SV40
Rt-PA
Zhang et al., 2008
L. japonica
Stable
Particle bombardment
SV40
bar
Zhang et al., 2008
Undaria pinnatifida
Transient
Particle bombardment
CaMV 35S
GUS
Qin et al., 1998
U. pinnatifida
Transient
Particle bombardment
SV40
GUS
Yu et al., 2002
Ulva pertusa
Transient
Electroporation
CaMV 35S,
GUS
Huang et al., 1996
U. pertusa
Transient
Particle bombardment
UprbcS
EGFP
Kakinuma et al., 2009
Brown
Note: AmCFP = humanized cyan fluorescent protein; AMT = Amino methyl transferase; CaMV 35S = cauliflower mosaic virus 35S promoter; CAT
= chloramphenicol acetyltransferase; EGFP = enhanced green fluorescent protein; FCP = fucoxanthin chlorophyll a/c- binding protein; GUS =
glucuronidase; HBsAg = human hepatitis B surface antigen; lacZ = bacterial beta-galactosidase; PtHSP70 = Porphyra tenera promoter; PyAct1 =
P. yezoensis actin 1 promoter; PyGAPDH = P. yezoensis glyceraldehyde-3-phosphate dehydrogenase; PyGUS = P. yezoensis glucuronidase; Rt-PA =
recombinant tissue plasminogen activator; sGFP = superfolder green fluorescent protein; S65T = mutated threonine; SV40 = a promoter; UBI =
ubiquitin (as gene promoter); UprbcS = Ulva pertusa ribulose-1,5-bisphosphate carboxylase/oxygenase (gene promoter); ZsGFP = humanized green
fluorescent protein; ZsYFP = humanized yellow fluorescent protein.
According to Mikami (2013), genetic transformation is reported in red and brown seaweeds
using the SV40 promoter; however, isolation of transgenic clone lines produced from distinct
single transformed cells, which is the final goal of the genetic transformation of seaweeds as a
tool, has not been reported, and seaweed genetic transformation is thus not fully developed.
Due to the problems with efficient genetic transformation systems, the molecular biological
studies of seaweeds are currently progressing more slowly than are the studies of land green
plants. Since a genetic transformation system allows the performance of genetic analysis of gene
function via inactivation and knock-down of gene expression by RNAi and antisense RNA suppression, its establishment will enhance both biological understanding and genetical engineering for the sustainable production of seaweeds and also for the use of seaweeds as bioreactors.
Though in vitro culture techniques as described above are currently being developed for seaweeds, which can create new genetic variants or promote clonal propagation in photobioreactors for high-end applications, most commercial seaweed cultivation, especially in the subtropical to tropical waters, is currently based on simple vegetative propagation because of economic
and farming advantages.
35
3.
MAJOR PROBLEMS OF FARMING SEAWEEDS
3.1
Disease and epiphytism
When a seaweed is suffering, we call it diseased, i.e. it is “dis-ease”. A seaweed is diseased when
it is continuously disturbed by some causal agents that results in an abnormal physiological
process that it disrupts its normal structure, growth, function or other activities. The concepts
of disease are the following (Singh, 2007): (i) the normal physiological functions of seaweed are
disturbed when they are affected by pathogenic living organisms and/or by some environmental
factors; (ii) initially, seaweed reacts to the “disease” causing agents, particularly in the site of
infection; (iii) later, the reaction becomes more widespread and histological changes take place;
(iv) such changes are expressed as different types of symptoms of the disease which can be visualized macroscopically; and (v) as a result of the disease, seaweed growth is reduced, deformed
or even dies.
Disease occurrence is generally driven by the interactions of three factors (Agrios, 2005; Garret et
al., 2009): (i) a susceptible host population; (ii) presence of a competent endophyte/malaise; and
(iii) a conducive (biotic and abiotic) environment (Figure 14).
FIGURE 14.
Infection triangle
e/
Su
sce
yth
pt
ph
ibl
do
eh
en
os
nt
t
ule
Vir
ma
lai
se
Infection
triangle
Conductive environment
Despite a developed technology of farming a seaweed, disease occurs, especially in areas where
stocking is intensive. Table 14 shows a summary of diseases caused by bacteria, fungi and epiphytes among farmed seaweeds.
36
TABLE 14.
Summary of seaweed diseases and epiphytism
Species
Disease name Causative organism(s)
Symptoms/effects
References
Chondrus crispus
Fungal parasite
Fungal parasite (Petersenia pollagaster)
Cavities and holes in fronds
Craigie & Correa, 1996
C. crispus
green spot or
green rot
Pathogen Lautitia danica
Infecting both cystocarpic and tetrasporangial region
Wilson & Knoyle, 1961; Schatz, 1984;
Stanley, 1992
C. crispus
Endophyte Acrochaete heteroclada and
A. operculata
Disrupts the cortica tissue of the host, slowing growth and
decreasing the capacity for regeneration
Gracilaria chilensis
Endophytic amoeba
Whitening, thallus decay and fragmentation
Red
White canopy
disease or
colourless
disease
Unknown, though probably similar to “ice-ice” in
K. alvarezii
G. heteroclada
Red spots
Agar-digesting bacteria/Vibrio sp.
Kappapphycus
alvarezii/ K. striatum
Ice-ice
Pseudomonas, Flavobacterium and
Actinobacterium
Eucheuma
denticulatum
Ice-ice
Marine-derived fungi (complex)
G. tenuistipitata
Correa & McLachlan, 1991, 1992, 1994;
Bouarab et al., 1999, 2001; Potin .,
1999, 2002; Brown ., 2003; Weinberger
et al., 2005
Correa & Flores, 1995; Buschmann et
al., 2001
Phap & Thuan, 2002
37
White to pinkish discolouration and gradual disintegration of the
thallus
Slow growth and greening of tissue
Whitening of thallus; softening of the branches or parts of
branches; development of white spots of dead tissue; and thallus
fragmentation
Lavilla-Pitogo, 1992
Uyenco et al., 1977; Largo et al., 1995a,
1995b, 1999
Solis et al., 2010
E.denticulatum
Penicillium waksmanii
Dewey et al.,1983
E.denticulatum
Scopulariopsis brevicaulis
Dewey et al., 1984
Kappapphycus
alvarezii/ K. striatus
Endophytic
filamentous
algae (EFA)
Neosiphonia saavatierii/red filamentous algae
Black goosebumps; presence of fine filamentous red algae; thallus
fragmentation
K.alvarezii/ K.
striatus
Endophyte
Colaconema infestans
Red endophytic filaments; alters the morphology and cellular
organization breakdown of cell wall
Critchley et al., 2004; Hurtado
& Critchley, 2006; Hurtado et al., 2006;
Vairappan, 2006; Vairappan et al., 2008;
Liu et al., 2009; Pang et al., 2011,
2012, 2015; Ateweberhan et
al., 2015
Araujo et al., 2014
K.alvarezii/ K.
striatus
Polysiphonia sp.
Palmaria palmata
Copepods (Thalestris rhodymeniae)
Galls or pinholes
Flavobacterium sp., Pseudoalteromonas sp., Vibrio
sp./Gram-negative bacteria
Lesions with wide green borders; slimy rots and holes in the blade
Pyropia yezoensis
Green-spot
disease
Tsiresy et al., 2016
Apt, 1988; Park et al., 1990
Nakao et al., 1972
38
P. yezoensis
Olpidiopsis
disease
Bleached portions on the blades; appearance of greenish lesions;
formation of numerous holes, followed by disintegration of the
entire blade
Olpidiopsis pyropiae/Oomycete
P. yezoensis
Diatom felt
Fragellaria sp., Licmophora abellata, Melosira sp.,
Navicula sp./Bacillariophyceae
P. yezoensis
Red-rot disease
Pythium porphyrae/Oomycete
P. yezoensis
Cyanobacter
ia felt
Filamentous and coccoid blue-green algae/
cyanobacteria
Dirty surface of blade; lesions and holes in the blade
P. yezoensis
White spot
disease
Phoma sp./Coelomycete
Bleaching of oyster shell with shell- boringconchocelis
Tsukidate, 1971; 1977
P. yezoensis
Suminori disease
Flavobacterium sp.
Pigmented
marine bacteria
Pseudoalteromonas
Populate the surface, preventing the colonization of other
seaweeds and invertebrate larvae
Egan et al., 2001
Hollowing of
stipes; stipe
blotch disease
Amphipod Amphitholina cuniculus; ascomycete
Phycomelaina laminariae
Boring of stipes and produces hollow
Myers, 1974; Chess, 1993
Unidentified parasitic micro-organism; amphipod
P. humeralis
Boring of stipes and produces hollow
Rheinheimer, 1992; Chess,
1993
Ascomycete Phycomelaina laminariae
Hyphae of P. laminariae penetrate the surface, leading to necrotic
tissue and reduced overall performance
S. digitata
Ascomycete Ophiobolus laminariae
Blackened patches of stipes
Sutherland, 1915
S. digitata
Ascomycete Petersenia sp.
Damages the stipes
Kohlmeyer, 1968
Klochkova et al., 2016
Dirty surface of blade; bleaching of blade
Red patches on the blade; blade’s colour changes from natural
brown, red to violet-red formation of numerous holes, followed by
disintegration of the blade
Ding & Ma, 2005
Green
Ulva lactuca
Brown
Alaria esculenta
Macrocystis pyrifera
Saccharina digitata
Black rot;
hollowing of
stipes
Stipe blotch
disease
S. digitata
Unknown hyphomycete
S. digitata
Endophyte Entocladia viridis
Nielsen, 1979
S. digitata
Endophyte Laminariocolax tomentosoides
Pedersen, 1976; Burkhardt & Peters,
1998
S. digitata
Endophyte Laminariocolax tomentosoides spp.
deformans
Galls and stipe coiling
Peters, 2003
S. digitata
Endophyte Laminariocolax aecidioides
Host thalli becoming thicker and stiffer, lowering their market
value
Peters, 2003
Bacterial flora (Flavobacterium/Cytophaga)
Lytic action on the viable cells
Ezura et al., 1988
S. japonica
Marine bacterium (Pseudoalteromonas
bacteriolytica)
Unique bacteriolytic activity and that induces damages
Yumoto et al., 1989a; Yumoto
et al., 1989b
S. japonica
Proteobacteria like Alteromonas, Vibrio
Detachment of gametophytes and young sporophytes from the
ropes
Ezura et al., 1988, Yamada et al., 1990
Pseudoalteromonas and Pseudomonas
Marginal portions of the diseased fronds turned greenish, become
soft, decay and disintegrate
Tang et al., 2001; Liu et al., 2002
S. japonica
Red spots
Causes contortion of the blade and blackening of the stipe
Kohlmeyer, 1968
Green rot
S. japonica
White rot
S. japonica
Malformatio n
disease
Sulfate-reducing bacteria
(Micrococcus)
Plasmolyzed oogonial and abnormal, malformed sporelings, which
subsequently die and drop off the cultivation lines
Falling-off
disease
Alginic decomposing bacteria
(Pseudomonas)
Sporelings falling off from the seeding ropes, especially during
summer
Chen et al., 1979
Frond-twist
disease
Polymorphic mycoplasma-like organism, (coccoid,
ovoid dumbbell, amoeboid shape)
Subnormally twisted fronds with great swollen stipes and very
shortened rhizoidal holdfast
Wang et al., 1983; Wu et al., 983;
Tsukidate, 1991
Hollowing of
stipes
Amphipod Ceinina japonica
Boring of stipes and produces hollow
Akaike et al., 2002
Ascomycete Phycomelaina laminariae
Hyphae of P. laminariae penetrate the surface, leading to necrotic
tissue and reduced overall performance
39
S. japonica
S. japonica
S. japonica
S. japonica
S. latissima
Stipe blotch
disease
Same course of development in green rot, only the fronds turn
white due to strong sunlight, high water temperature and lack of
nutrients
Andrews, 1976
Wu et al.,1983
S. latissima
Endophyte Entocladia viridis
Nielsen, 1979
S. latissima
Endophyte Laminariocolax tomentosoides
Lund, 1959
S. latissima
Endophyte Laminariocolax aecidioides
Host thalli becoming thicker and stiffer, lowering their market
value
Peters & Ellertsdottir, 1996; Heesch &
Peters, 1999; Peters, 2003
Undaria pinnatifida
Spot rotting
U. pinnatifida
Shot-hole
disease
Vibrio
Brown spots appearing on the thallus blade
near the midrib, which subsequently fuse together and spread
onto the pinnate part of the blade
U. pinnatifida
Green spot
disease/rot
Unspecified bacteria
Small holes with green margins
U. pinnatifida
Green decay
disease
Aeromonas, Flavobacterium, Moraxella,
Pseudomonas and Vibrio
Kimura, et al., 1976
Vibrio logei
Tsukidate, 1991
Ishikawa & Saga, 1989; Vairappan et al.,
2001; Kang, 1982
Jiang et al., 1997
40
U. pinnatifida
Yellow hole
disease
Unspecified bacteria
U. pinnatifida
Spot rotting
Unspecified bacteria
Kito et al., 1976
U. pinnatifida
Spot decay
Bacterium Halomonas venusta
Ma et al., 1997a, 1997b, 1998
U. pinnatifida
Pin hole
Frond-mining nauplii of harpacticoid copepod
(Amenophia orientalis, Parathalestris infestus,
Scutellidium sp. and Thalestris sp.)
Tsukidate, 1991; Ho & Hong, 1988; Rho
et al.., 1993
U. pinnatifida
Tunnel
Gammeride amphipod, Ceinina japonica
U. pinnatifida
Chytrid blight
Oomycete, Olpidiopsis
The fungus affects sporophytes, where it grows inside host cells,
killing them slowly
Tsukidate, 1991
U. pinnatifida
Endophytic
brown alga
Laminariocolax aecidioides
Host thalli becoming thicker and stiffer, lowering their market
value
Akiyama, 1977; Yoshida & Akiyama,
1978
Small holes with yellow margins
Invades the midrib of U. pinnatifida through the holdfast and
bores a tunnel, which may cause the longitudinal separation of
the entire frond through the midrib
Ishikawa & Saga,
1989; Vairappan et al., 2001; Vairappan
et al., 2001
Kang, 1982
3.2
Social and financial
Issues on social problems pertinent to seaweed farming stem from the unacceptability by the
community to the introduction of a novel farming system. This is brought on mainly if such
farming system affects the immediate environment.
One of the biggest problems of seaweed-carrageenan farming is the accessibility to financial
assistance, especially in areas where cyclones or typhoons occur, such as the Philippines. Normally, farming structures and propagules are destroyed when the typhoon signal is No. 2 or
higher. The capacity to rehabilitate is a major problem. The need to have crop insurance in
seaweed aquaculture activity is important so that in times of calamities seaweed farmers can
claim a certain amount of the lost crop and structures to restart farming.
4.
IMPACT OF SEAWEED FARMING
4.1
Socio-economic impact
The comprehensive report of Valderamma et al. (2013), which includes six case studies of carrageenan seaweed farming in six different countries (India, Indonesia, Mexico, the Philippines,
Solomon Islands and the United Republic of Tanzania), attests to the economic benefits of Kappaphycus farming in the tropics and subtropics. In the temperate countries, reports include an
economic analysis of Laminaria digitata farming in Ireland by Edwards and Watson (2011); a cost
analysis for ethanol produced from farmed seaweeds by Philippsen et al. (2014); a new bioeconomy for Norway by SINTEF (2014); and economic feasibility of offshore seaweed production in
the North Sea by Van den Burg et al. (2013). All these reports clearly show that seaweed farming
is economically beneficial to farmers in particular and the local and national economy in general.
4.2
Ecological-environmental impact
Seaweed farming is an extractive aquaculture whose very process of production of valuable biomass renders the sea’s various ecosystem services with ecological and economic values
(Chopin et al., 2008, 2010; Neori et al., 2007; Radulovich et al., 2015). Seaweed farming adds
oxygen during photosynthesis and cleans seawater from excess nutrients (nitrogen, phosphorus
and others). Nutrient extraction, or uptake, cleans water effectively and thoroughly through a
process known as bioremediation (Forster, 2008). Seaweed farming enhances biodiversity and
fisheries (Radulovich et al., 2015). Seaweeds are carbon sinks that can reduce ocean acidification
through uptake of CO2 from water.
Shading and alkalization may harm and benefit different local biological activities, competing
with phytoplankton and therefore filter feeders, but at the same time aiding calcification of
shellfish and corals, which suffer from ocean acidification (Branch et al., 2013).
Among the red seaweeds being farmed, Kappaphycus is drawing much attention in places where
it is being introduced. The literature shows that this seaweed is endemic in the tropics such as
Indonesia, Malaysia and the Philippines; its first successful commercial farming was reported in
the Philippines in the early 1970s (Doty, 1973; Parker, 1974; Doty and Alvarez, 1981). Since then,
it has been introduced in almost 30 countries worldwide. Such introduction without prior scientific and quarantine protocols and proper management led to some negative impacts in Hawaii,
United States of America, as claimed by Rodgers and Cox (1999), Smith et al., (2002) and Conklin
and Smith (2005), and in India by Chandrasekaran et al. (2008), instead of bringing economic
benefits to coastal families. However, the latest report on such bioinvasion and coral encroach-
41
ment was negated by the report of Mandal et al., (2010) for the following reasons: (i) lack of a
functional reproductive cycle; (ii) low spore viability; and (iii) the absence of microscopic phases
in the life cycle of Kappaphycus, coupled with the abundant presence of herbivores, restricted
the further spread of this alga.
5.
DRIVERS OR MOTIVATIONS TO PURSUE OR EXPAND
FARMING
The expansion or increase in seaweed farming in terms of production is mainly due to increasing demand for food, feed (animal) and, more recently, fuel. The global demand for seaweed
biomass is rising. Large companies using algae in their products require a regular and reliable supply of the material, both in quantity and quality. Western Europe and elsewhere will
continue to improve farming techniques to increase production, mainly because of the high
market value of the different products derived from seaweeds (Holdt, 2011). Figure 15 shows
the pyramid of the seaweed product markets.
FIGURE 15.
Pyramid schematic of seaweed product markets
Pharmaceutical / Cosmetics
Market value
100€/Kg
Fine chemical / food
10€/Kg
Protein (feed)
1€/Kg
Biomass for
energetical use
Market size
5.1
Food
Asians will continue to consume seaweeds as part of their daily diet. There is a rising awareness
of health and nutritional benefits from seaweeds in the Western countries. Likewise, there is
a growing use by food processors in new applications that include seaweed pasta, mustard,
rillettes and pâtés. Also, there is a high demand from the catering and food service sector that
requires seaweed recipes. Hence, cultivation of economically important seaweed will expand as
the population grows.
5.2
Feed (aquaculture)
The commercialization of land- and sea-based IMTA in western Europe will open more opportunities to an immense use of seaweed as part of the diet of fish such as salmon, rainbow
trout, cod, sea bass and other high-value fish. This is simply because several earlier studies have
42
demonstrated the positive effects not only in terms of the increased growth rate, but more
importantly, on the prevention of diseases (Wan et al., 2016; Walker et al., 2009; Valente et al.,
2006). Likewise, hogs fed with seaweed resulted in more milk of the sow, decreased mortality
by 50 percent, cut the use of antibiotics by 50 percent, improved health management, reduced
feed intake (gut health), made hogs ready for slaughter two to three weeks earlier, improved
taste (industrial taste panel), and doubled omega-3 in pork (Kraan, 2015). The high demand of
seaweed-fed abalone will continue, as the growing population prefers traceable marine food.
The newly emerged application of seaweed in the shrimp diet will be developed and refined
further. For these reasons, responsible and sustainable farming of seaweed will increase in the
next few years.
5.3
Fuel
Traditionally, seaweeds have not been considered as feedstock for bioenergy production,
but have been used in food, in medicine or as fertilizer, and in the processing of phycolloids
and chemicals (Bixler and Porse, 2011). The cultivation of algal biomass for the production of
third-generation biofuels has received increasing attention in recent years, as seaweeds can be
produced in the marine environment and on non-arable lands. Production yields of algae per
unit area are significantly higher than those for terrestrial biomass (Wei et al., 2013; Schenk
et al., 2008). The chemical composition of algae makes it suitable for conversion into biofuels,
especially the subtidal large brown kelps of the order Laminariales (Hughes et al., 2013) and
Ulva (Bruton et al., 2009).
Seaweeds are already farmed on a large scale in Asia and to a lesser extent in Europe, primarily
in France, and on a research scale in Scotland (Kelly and Dworjanyn, 2008). Western Europe,
Ireland in particular, is becoming aggressive in research and development for a marine bioenergy
and biofuel industry (Roberts and Upham, 2012). Biofuel production from macro-algae is in its
infancy. There is a strong collaboration in the private sector, such as Statoil ASA, which entered
into a partnership with Seaweed Energy Solutions AS (SES) and Bio Architecture Lab (BAL) to
develop a macroalgae-to- ethanol system in Norway. The aim of the partnership is to develop a
10 000 ha seaweed farm off the coast of Norway, which will produce 200 000 tonnes of ethanol
(equivalent to 2 percent of the European Union’s ethanol market) (Ystanes and Fougner, 2012).
SES is developing the technology for large-scale cultivation and harvesting technology, while
BAL is responsible for developing the technology and the process to convert the macro-algae
into ethanol (Murphy et al., 2013).
Though several preliminary investigations have been conducted to assess the technical feasibility, environmental viability and economic profitability of seaweed farming fo fuel (Watson,
2014; Valderamma et al., 2013; Watson et al., 2012), numerous parameters (such as method
of cultivation, species of seaweed, yields of seaweed per hectare, time of harvest, method of
harvesting, suitability of seaweed to ensiling the gross and net energy yields in biogas, carbon
balance, cost of the harvested seaweed, and cost of the produced biofuel) have to be developed
economically to obtain viable algae biofuel production.
6.
CONSERVATION AND SUSTAINABLE STRATEGIES
Conservation is a careful preservation and protection of resources that includes a well-planned
management of said natural resource to prevent exploitation, destruction or neglect. There is
biodiversity of seaweeds within species (genus), between species, and of ecosystems; hence, each
43
species has its own peculiar characteristics to adapt in a certain habitat. Seaweeds, both harvested and farmed, are important sources of livelihood to humans. Conserving and sustaining
these resources for the benefit of mankind are imperative.
A sustainable livelihood is one that can be carried out over the foreseeable future without
depleting the resources it depends upon and without depriving others of a livelihood. In order
for a livelihood to be sustainable, there should be: (i) economic development; (ii) social equity;
and (iii) environmental protection. Sustainable development can be achieved if decisions are
made to be economically profitable, biologically appropriate and socially acceptable (Figure 16)
(Eigner-Thiel et al., 2013) (Circular Ecology, 2016).
FIGURE 16.
Sustainability paradigm
Social
Equitable
Bearable
Sustainable
Economic
Sustainability
Viable
Environmental
Sustainability
Curtesy circularecology.com
Currently, with intensive fed aquaculture (finfish and shrimp) throughout the world is rapidly
increasing, environmental impact is the main concern. A large amount of this concern pertains to
the direct discharge of significant nutrient loads into coastal waters from open waters and with
the effluents from land-based systems. The only way to mitigate this environmental concern
is to adopt an aquaculture system that is sustainable and balanced, a system known as integrated multi-trophic aquaculture (IMTA) (Chopin et al., 2001). Aquaculture is the world’s fastest
growing food production sector, and is associated with environmental, economic and societal
issues. IMTA offers an innovative solution for environmental sustainability, economic stability,
and societal acceptability of aquaculture by taking an ecosystem-based management approach.
IMTA is the farming, in proximity, of aquaculture species from different trophic levels and with
complementary ecosystem functions, so that one species’ excess nutrients are recaptured by the
other crops and synergistic interactions among species occur (Chopin et al., 2013). By integrating
fed aquaculture (finfish, shrimp) with inorganic and organic extractive aquaculture (seaweed
and shellfish), the wastes of one resource user becomes a resource (fertilizer or food) for the
44
others. Such a balanced ecosystem approach provides nutrient bioremediation capability, mutual
benefits to the co-cultured organisms, economic diversification by producing other value-added
marine crops, and increased profitability per cultivation unit for the aquaculture industry.
In order for seaweed farming to be sustainable elsewhere, the following are to be implemented:
(i) expansion of farming areas, wherever possible and profitable, and subject to the needs of
other sectors and of environmental health; (ii) improvements in productivity through the development and wide adoption of better aquaculture practices, to include improved quality of seed
supply, establishment of land-sea based nurseries, including innovative approaches such as IMTA;
(iii) increased investment in research, development and extension (RD&E) to meet expected challenges, including disease risks, climate change and introductions of non-indigenous species; and
(iv) strong collaboration among government agencies, academia and the private sector. Table 15
presents the conservation and sustainability strategies for farmed seaweeds.
TABLE 15.
Conservation and sustainable strategies for farmed seaweeds
Conservation
and sustainable
strategies
Capacity enhancement
of human resources
Action plans
• Active enhancement of public promotion and environmental education through regular training/
workshops/seminars
• Cross-country/area visits to successful seaweed areas/farmers
• National and international collaboration and networking
• Improve scientific knowledge and strong cooperation with local and international societies and
stakeholders working on the conservation of marine resources
Diversified livelihood
Sound ecosystembased management
Secured sustainability
• Introduction of invertebrate aquaculture and sea-ranching, such as sea urchins, sea cucumbers and
sea abalone and other high-value animals, instead of fisheries/capture in areas where there is natural
population
• Cultivation of other economically important seaweeds with bioactive, biofuel, pharmaceutical,
cosmetic and nutraceutical potential
• Adaption of better aquaculture practices
> Sufficient buffer space between lines and farms to allow free water movement
> Reduction of the number of farms in dense cultivation areas to include maximum carrying capacity
> Use of appropriate cultivation method suitable to the environmental conditions of a given area
> Use of biodegradable planting materials
• Proper zoning of aquaculture activities
• Adaption of a no-no policy of placing seaweed farms near or on top of coral reefs or in marine
protected areas
• Prevention of marine pollution coming from inland domestic and industrial effluents and sea-oil
pollution
• Large-scale production
> Production on a large scale in order to secure profitability, stable operation of the production
facilities, and build up a buyer’s market
> Maximizing the potential of macro-algae using the biorefinery approach
• Products
> Development of other product applications of agarophytes, carrageenophytes, alginophytes and
some green macro-algae
> Development of biorefinery processes, which make possible parallel utilization of several
components (pharmaceuticals and cosmetics, food and feed, bioplastic and polymers, bulk
chemicals and fuel, and heat and energy)
> Development and testing of animal feed based on seaweed biomass
> Securing marketing channels and maturing of the market for seaweed and products based on
seaweed
> Strong cooperation between industry, academia/research centres and government authorities
45
7.
ENHANCEMENT PROGRAMME
7.1
Education
Development of human resources through scholarships and fellowships is encouraged, especially
in developing countries, to pursue professional and personal advancement in the different fields
of specialization in seaweeds for graduate and post-graduate programmes. Such education will
prepare students to embark in tougher responsibilities needed in the community and the industry. A number of scholarships are being offered by developed countries, such as Australia, mainland European countries, Japan, United Kingdom of Great Britain and Northern Ireland, and the
United States of America, and are highly competitive.
7.2
Research and training
Skills training is designed both to improve student effectiveness as researchers and to equip
them with the skills they will need in a career after graduating – whether to choose to follow
an academic or a non-academic career path. The structure and design of Ph.D. programmes
should incorporate generic skills and be formulated with direct engagement with employers and
enterprises where appropriate. Training helps people improve their competencies, which leads
to better performance appraisals.
Worldwide, state universities and colleges as well as research centres have good programmes
for seaweed research and training. Students and trainees are given the opportunity to conduct
research according to the needs of the industry under the supervision of a professor or a scientist. They are trained to: (i) conceptualize and write a proposal; (ii) conduct the study with little
supervision; (iii) collect, analyse and interpret the data; (iv) make conclusions; (v) write a manuscript for publication; and (vi) share the results to the scientific community through attendance
at symposia and congresses.
It is in the stage of research and training that individuals will establish a strong working relationship with their mentor, peers, the private sector and community.
8.
ROLE OF INTERNATIONAL AND REGIONAL ASSOCIATIONS
IN THE DEVELOPMENT AND MANAGEMENT OF FARMED
SEAWEEDS
There are several international and regional associations that are involved in the development
and management of farmed seaweeds, as shown in Table 16. These associations have different
mandates to fulfil for the betterment of the community and industry.
46
TABLE 16.
International, regional and local associations, organizations and societies engaged in seaweed
research and other related activities
Location
Name of
organization/
society
Objectives
Asia-Pacific
Asian Pacific
Phycological
Association
• Develops phycology in the Asia-Pacific region, to serve as a venue for the exchange
of information related to phycology and to promote international cooperation among
phycologists and phycological societies in the Asia-Pacific region.
• Holds meetings at least once every three years.Asia-Pacific
Asia-Pacific
Asia-Pacific Society
for Applied Phycology
• Cooperates with national and international phycological organizations.
Australia
Australasian Society
for Phycology and
Aquatic Botany
• Promotes, develops and assists the study of, or an interest, in Australia
phycology and aquatic botany within Australasia and elsewhere.
• Establishes and maintains communication with people interested in phycology and
botany.
China
China Algae Industry
Association
• Promotes the rationalization of alga, producing and processing product mix,
management system and business organization.
• Contributes to the alliance of industry, agriculture and business.
• Coordinates the relation of production, supplement and marketing.
China
China Phycological
Society
• Builds China’s largest professional information service platform, science and
technology innovation platform, and brand promotion platform for the algae industry.
China - Taiwan
Province
Taiwanese
Phycological Society
• Enhances and strengthens algal academic research.
• Promotes algal awareness and develops algal applications
Europe
British Phycological
Society
• Advances education by the encouragement and pursuit of all aspects of the study
of algae, and publishes the results of the research in a journal as well as in other
publications.
• Publishes the British Journal of Phycology and the newsletter, The Phycologist.
Europe
Federation of
European Phycological
Societies
• Provides a forum for all European phycological societies and individuals with an
interest in phycology; enables, promotes and enhances algal (including cyanobacterial)
research, education and other activities; increases public awareness of the importance
of algae and cyanobacteria; and contributes to public debate and policy issues
involving these organisms throughout Europe.
Europe
Hellenic Phycological
Society
• Promotes basic and applied phycological research, organizes congresses, and develops
international relationships.
Indonesia
Asosiasi Rumput Laut
Indonesia
• Develops downstream seaweed industries to create more added value from this
marine commodity and to create job opportunities.
Japan
Japanese Society of
Phycology
• Promotes research that is related to algae and phycology, and serves as a central hub
of people who are interested in phycology.
Republic of Korea
Korean Society of
Phycology
• Promotes publications of algae, which deal with phylogenetics, taxonomy ecology
and population biology, physiology and biochemistry, cell and molecular biology, and
biotechnology and applied phycology.
• Publishes the journal Algae.
Philippines
Philippine Phycological
Society, Inc.
• Promotes the science of phycology in the Philippines.
Philippines
Seaweed Industry
Association of the
Philippines
• Develops better technology for growing and processing better quality colloids in
alliance with academic institutions and international associations.
South America
Brazilian Society of
Phycology
• Gathers together people and institutions interested in the development of phycology.
• Promotes and stimulates teaching and research on algae and other photosynthetic
aquatic organisms.
South America
Chilean Phycological
Society
• Promotes phycological research, and the development, scientific knowledge and
protection of the phycological flora in Chile.
Southeast Asia
ASEAN Seaweed
Industry Club
• Promotes strong cooperation and networking among the ASEAN countries.
• A forum of national and foreign professionals interested in the world of algae.
47
Location
Name of
organization/
society
Objectives
Spain
Spain Phycological
Society (Sociedad
Española de Ficologia)
• Establishes partnerships between phycologists, public and private research
organizations, and companies interested in the study and applications of algae.
USA
International
Phycological Society
• Develops phycology distribute phycological information; cooperates among
international phycologists and phycological organizations; and convenes the
International Phycological Congress every four years.
USA
International Seaweed
Association
• Convenes the International Seaweed Symposium every three years, the leading global
forum for researchers, industrial companies and regulators involved in the seaweed
sector.
USA
International Society
for Applied Phycology
• Promotes research, preservation of algal genotypes and the dissemination of
knowledge concerning the utilization of algae.
USA
Marinalg International
• Promotes the image and uses of seaweed-derived hydrocoloids in food,
pharmaceuticals and cosmetics.
USA
Phycological Society of • Promotes research and teaching in all fields of phycology; publishing the Journal
America
of Phychology
9.
SOURCES OF DATABASES
9.1
Regional and international centres
Only a few countries and regions have their own seaweed centres that cater to the needs of the
industry and community. The Western countries have centres dedicated mainly for basic and
applied research on algae that may be absent in the developing countries. However, a small
research laboratory is normally present in the university or in fisheries institutions. Table 17 lists
international centres that have strong collaboration with other institutions/academia or industry
in and out of the region with their respective mandates.
TABLE 17.
Some international algae centres
Name and website
University/private sector
Mandate
AlgeCenter Danmark
(www.algecenterdanmark.dk)
Aarhus University; Kattegatcentret;
Danish Technological Institute
Research in the areas of: (i) biorefinery; (ii)
algae growing; and (iii) energy production
Centre d’Etude et de Valorisation des
Algues (CEVA) (www.ceva.fr)
Pleubian, France
Dedicated to the study and enhancement
of algae (macro and micro) marine plants
and marine biotechnology
MACRO – the Centre for Macroalgal
Resources & Biotechnology (https://
research.jcu.edu.au/macro)
James Cook University, Australia
Develops and commercializes marine and
freshwater macro-algae for fuel, feed and
fertilizer applications
Norwegian Seaweed Technology
Center
(www.sintef.no)
SINTEF Fisheries and Aquaculture; SINTEF
Materials and Chemistry; Norwegian
University of Science and Technology
(NTNU); Department of Biology;
Department of Biotechnology
Develops technology within industrial
cultivation, harvesting, processing and
application of seaweed in Norway
Seaweed Energy Solutions AS
Norway, Portugal and Denmark
Focuses on large-scale cultivation (www.
seaweedenergysolutions.com) of seaweed
primarily for energy purposes
48
The biggest storage of seaweed information in terms of taxonomy, description and distribution
is found in www.algaebase.com. All universities and research institutions that have seaweed
programmes have an herbarium of their local species, as well as algae journals and books in their
libraries.
9.2
Dissemination, networking and linkages
Scientific knowledge coming from research can be disseminated through the following ways: (i)
publication in peer-reviewed journals, symposium proceedings and books; (ii) presentation of
results in different symposia and congresses; and (iii) writing in popularized magazines, newletters, brochures and flyers for the industry.
Networking is important in the seaweed community. There is a need to work together to develop
seagriculture, or sea farming, in order to cater to the needs of the industry. Vertically integrated
supply chains require a lot of energy from small companies. There is a need to improve the value
chain for
better efficiency and maximize shared benefits among the seaweed community. There are
mutual benefits and assistance derived from linkages and networking activities with both local
and international organizations. Linkages and networking are different in the degree of commitment by the partners. In linkages, the relationship between partner organizations is quite
loose. It intends to serve the members of both sides according to their respective needs, interests
and objectives. It creates bonds together to solicit support and assistance for purposeful activities.
Networking, on the other hand, is much stronger, usually because the groups and agencies have
common objectives and beneficiaries. Networking is basically extending the outreach of the
resources in different ways so as to increase the effectiveness of the programme. The areas of
operation can also be increased through networking. A network is composed of several institutions, universities or research centres that bind together for a common goal. They work
together to attain common objectives, undertake innovative practices, and update members
regarding breakthroughs in different disciplines. Table 18 lists some of the active networks in
different regions.
TABLE 18.
Various networks involved in seaweed farming and allied activities
Network
Objectives
Asian Network for Using Algae
as CO2 Sink
Encourages collaboration among member countries in conducting research in sustainable CO2
removal by marine-life mechanisms.
Canadian Integrated
Multi-Trophic Aquaculture
Network
Provides interdisciplinary research and development and highly qualified personnel training in the
following linked areas: (i) ecological design, ecosystem interactions and biomitigative efficiency; (ii)
system innovation and engineering; (iii) economic viability and societal acceptance; and (iv) regulatory
science.
Danish Seaweed Network
Promotes the production, application, communication and knowledge of seaweed, and also to
strengthen the national collaboration.
Global Seaweed Network
Develops a programme, which over the next 5–10 years will enhance and develop the global
seaweed community into an internationally recognized and respected scientific body that can
innovate, provide knowledge and tools for scientific research, aquaculture, conservation and society,
influence policy-makers and enable economic progress.
49
Network
Objectives
Netalgae (France, Ireland,
Norway, Portugal, Spain,
United Kingdom)
Creates a European network of relevant stakeholders within the marine macro-algae sector. Compiles
information from different regions that will result in a wide-ranging policy study of existing practices
within the macro-algae industry.
Analyses the results that will establish a best practice model and suggests policies for the successful
sustainable commercial utilization of marine macro-algae resources.
Nordic Algae Network
(Denmark, Iceland, Norway,
Sweden)
Helps the partners to a leading position in the algae field for commercial utilization of high- value
products and energy from algae.
Increases the synergy and facilitates collaboration between partners.
Norwegian Latin American
Seaweed Network Norwegian
Seaweeds Network
Encourages cooperation among the seaweed stakeholders across Latin America and Europe in order
to support the development of the seaweed sector.
Strengthens interest and knowledge of benthic algal taxonomy, systematics and species
identification, and promotes collaboration and exchange of information.
REBENT
(France – national network
coordinated by IFREMER)
Collects and organizes data concerning marine habitats and benthic biological communities in the
coastal zone to provide relevant and coherent data to allow scientist administrators and the public to
better determine the existing conditions and detect spatiotemporal evolution.
10.
EXCHANGE PROGRAMMES
10.1 Information
Science and technology provide critical tools that help address national and global needs.
Freedom of scientific exchange and stronger scientific collaboration to benefit humankind is of
paramount importance. Open exchange of information and ideas is critical to scientific progress.
To achieve this end, there should be: (i) promotion of a strong, non-governmental, scientific
publishing enterprise that ensures access to information and exchange of scientific ideas and
information among all parties with legitimate uses while appropriately protecting copyright and
security-related information; (ii) assurance of the quality of science and technological advancement through open, rigorous and inclusive peer review scientific publishing; and (iii) open interactions among scientists, engineers and students from across the globe.
The discovery of computer technology has opened many opportunities to gain access to more
than one system to gather data or exchange information. Open access and exchange of information is one of the core values of academics; a computer system that limits access is frustrating
at best. Open access is part of the open science movement and covers various initiatives and
projects across the globe to make academic studies and results available to a wider readership.
Open access promotes knowledge transfer by mouse click. There are alternatives to expensive,
restricted access to academic publications, for example, PLOS ONE, the Public Library of Science’s
international online journals. The publications can be accessed from any computer with an Internet connection; the author retains the copyright; manuscripts are published relatively quickly;
they are peer reviewed by experts; quality and impact can be determined using post-publication
tools; and users can discuss the articles in communities. As open access publications are available free of charge throughout the world, even people in poorer countries who usually lack the
financial means can access and use them.
Regular members of the International Seaweed Association have free access to the Journal of
Applied Phycology, a journal that publishes articles on micro- and macro-algae (seaweeds) with
four issues each year.
50
10.2 Scientists and experts
Scientists and experts play crucial roles in the exploitation, management, conservation and sustainability of seaweed resources. Results of their scientific studies are used to formulate policies
for the government to adapt for implementation.
According to Dr Houde of the Chesapeake Biological Station, United States of America, scientists
have the difficult task of walking the fine line between traditional “science-worthy” science, or
making the news. Traditional science takes time, as the peer review process is typically a slow
one, even though it helps to minimize errors. Often, it moves too slowly for policy, which has
now begun to turn to “post-normal” science, which pools the collective advice of experts. On the
other hand, making the news often means bold and dramatic statements, which is sometimes
risky. Science-based decision- making is not that straightforward. One can use models and mathematical equations to predict various outcomes, but one cannot guarantee those results. Thus,
when real life does not follow model predictions, people lose faith in the science.
Seaweed farming is centred on the management of the environment and sustainability of the
commodities. It takes several years for scientists and experts to transfer the science-based technology to the industry. Trials of farming Kappaphycus and Eucheuma in the Philippines started in
1965 and it was only in 1971 when the first harvest of seaweed for export purposes was attained
(Doty and Alvarez, 1981). Also, the introduction of IMTA in Canadian waters started as early
as 2000 and became commercial several years after. Though biological and economic results
were positive, social acceptability was a critical component in aquaculture sustainability (Barrington et al., 2009). Scientists and experts, together with the different stakeholders, met several
times to discuss the importance and significance of IMTA. All agreed that IMTA has the potential
to reduce the environmental impacts of salmon farming, benefit community economies, and
improve industry competitiveness and sustainability. This successful aquaculture system is currently being replicated either on an experimental or near commercial stage in western Europe
(Holdt and Edwards, 2014; Lamprianidou et al., 2015; Freitas et al., 2016).
Scientific and technological development is impossible without efficient communication between
scientists or technologists and the community. Such that, a higher level of scientific research can
be achieved through collaboration.
10.3 Test plants
Only test plants preserved in silica gels and dried samples previously soaked in 10 percent formaldehyde and later drained are allowed to be sent by courier to other universities or institutions
outside from its point of origin for collaborative work. This is especially true in developing countries, which lack the facilities to analyse the samples for a specific test. The test plants serve as
the share of the collaborative study, and ultimately, part of the authorship when the results are
written and submitted to a peer- reviewed journal for possible publication. No fresh test plants
are allowed by courier for scientific study. However, live test plants are allowed by the scientist to
bring personally after proper documents from point of origin to final destination are in order. If
no prior agreement is made with the provider of test plants for research and scientific purposes,
a due recognition through acknowledgement at the end of the report or paper is appropriate.
51
11.
CONCLUSIONS
The farming of economically important seaweeds for food is dominantly done in Asia for the
past several decades and will continue to increase as population increases. On the other hand,
the farming of seaweed for feed and fuel purposes will be centred in the Western countries.
Also, people in Western countries are increasing their seaweed consumption as part of their diet
for health reasons.
China, Japan and the Republic of Korea are the leading producing countries of brown seaweeds
(Saccharina and Laminaria) and red seaweed (Pyropia), while Indonesia and the Philippines are
the top leading producers of Kappaphycus and Eucheuma. It is surprising to learn that Indonesia
has surpassed China and Chile in the production of Gracilaria starting around 2013. Indonesia is
presently the world’s number one producer of farmed red seaweeds, notably Eucheuma, Gracilaria and Kappaphycus.
Innovations in farming systems are being done because of disease and epiphytism problems
brought on by climate change. Seaweed farmers with the technical assistance of scientists and
experts will continue to work together for the improvement of crop management, productivity
and production. One example of a culture farming modification is the traditional farming of
Kappaphycus, which has now shifted from shallow waters to deeper waters to avoid elevated
surface water temperature that adversely affects productivity and production.
Use of plantlets from spores remains to be used in the lab for outplanting purposes with improvements in nutrition-temperature-light requirements. Although several successful studies were
reported on the regeneration of plantlets of Kappaphycus and Euchuema from callus through
micropropagation using different culture media, their economic viability in the field has yet to
be tested further, though initial trials have been started. Likewise, the use of seaweed extract
as a biostimulant in the micropropagation of Kappaphycus has proven successful and field trials
are in progress. At present, vegetative propagation still dominates the commercial farming of
Kappaphycus and Eucheuma. The successful hybridization of Saccharina japonica using gametophytes and sporophytes in China (Dongfang No. 7) may provide a model for domestication to
other brown seaweeds (kelp).
Currently, seaweed genetic transformation is not fully developed despite several studies reported.
Because a genetic transformation system would allow to perform genetic analysis of gene function via inactivation and knock-down of gene expression by RNAi and antisense RNA suppression, its establishment will enhance both our biological understanding and genetical engineering for the sustainable production of seaweeds and also for the use of seaweeds as bioreactors.
IMTA as a holistic aquaculture system has been tested to be technically feasible, environmentally friendly, economically viable and socially acceptable in the Western countries and China.
Its replication in other countries, especially in countries engaged in intensive shrimp and finfish
aquaculture, has yet to be introduced or developed.
Conservation and sustainability of farmed seaweeds are the ultimate goals to ensure that the
biomass needed for its final product is maintained commercially.
Seaweed international centres, societies, organizations and associations, and networking among
scientists and experts will continue to play important and significant roles in the further development and ultimate sustainability of farmed seaweeds, which are good for food, feed and fuel.
52
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