Yeallow Sea Fisheries Research Institute,

Chinese Academy of Fishery Sciences

Theory and technology of Pyropia culture in China

Wang, Wen Jun

Tel: +86-15066238219

E-mail: wjwang@ysfri.ac.cn

DIVISION OF THEORY AND TECHNOLOGY FOR MARICULTURE

 1. Background
1.1 Classification of Bangiaceae

147 (infra) species deposited in algaebase,

16 resolved[2]
Bangia

Dione
Sutherland et 5 species defined in Flora Algarum
Bangia Minerva
al. 2011 [1] Marinarum Sinicarum [3]
Pseudobangia
Bangia Lyngbye 1819
B
a 3 unnamed genera
n
g
i
Porphya 277 (infra)species deposited in algaebase, 59
a
accepted[2]
c
e Boreophyllum
a
e Clymene
21 species and 3 variations in
Fuscifolium
China [3]
Porphyra Sutherland et
al. 2011[1] Lysithea

Porphyra C. Agardh Miuraea

1824 81 (infra)species deposited in algaebase, 73
Pyropia
resolved[2]
Wildemania
1.2 Nutrient content of Pyropia and kinds of Pyropia food

1. High protein, essential amino acids (EAAs), delicious amino acids content, EAA/TAA and DAA/TAA[4].
2. Low lipid content but high ratio of unsaturated fatty acids (TUFAs) to total fatty acids (TFAs), with EPA accouting for up to 80% of TUFAs
in some species[4].
3. The following data in Fig. 1 come from a publication of Pyropia yezoensis[5] .
4. Kinds of nutritious and delicious foods have been made of Pyropia (Fig. 2).

6%
EAA/TAA: ~35%
Protein 38%
Lipid DAA/TAA: ~50%
Carbohydrate
TUFA/TFA: 50-70%
f
Ash
53%
EPA/TUFA: 60-80%
3%
Fig. 1 The main nutrient content of a Pyropia yezoensis strain[5].

a b g

d
c
h

e
Fig. 2 Kinds of foods made of Pyropia. a: Dried P. haitanensis and P. yezoensis; b, c, d, e, f, g, h: Soup, fried eggs, sauce, roll, flavored laver
broken, sandwich, and sushi made of/with laver.
1.3 Statistics of seaweed farming in China

1. Pyropia contains the highest culture area and Saccharina contains the highest annual yield (Fig.3).
2. The culture area of Pyropia has been increasing and doubled in the recent 10 years and becomes the highest among all the cultured seaweeds in
China [4].

a. Scale of cultivated seaweeds in China in 2017 (103 ha) b. Yield of cultivated seaweeds in China in 2017 (104 ton)

Saccharina
Undaria
Pyropia 17.3305 (8%)
44.236 (31.5%)
Gracilaria
Eucheuma
79.607 (56.6%) 148.6645
Sargassum
(68.8%)
The others

Fig. 3 Culture area (a) and annual yield (b) of the main cultured seaweeds in 2017 in China.
 1.4 Distribution of Pyropia (Porphyra) resource and industry in China

1. The wild Pyropia (Porphyra) resources are widely distributed along the coast
of China (Fig. 4, yellow line).
2. The farmed Pyropia industries are mostly distributed from Guangdong to
Shandong province (Fig. 4, red line) with the highest culture area in Jiangsu
province (Fig. 5a) and the highest yield in Fujian province (Fig. 5b).

a 50 47.255

Culture area in 2017（103

40

ha）
30

20 15.178
13.709

10
2.978
0.487
0
Shandong Jiangsu Zhejiang Fujian Guangdong

7 6.3509
b
Yield in 2017（104 ton）
6
5 4.186 4.2632
4
3
2 1.4931
1.0373
1
0
Shandong Jiangsu Zhejiang Fujian Guangdong

Fig. 4 Distribution of Pyropia (Porphyra) resource along the coast of China. Fig. 5 Culture area (a) and annual yield (b) of Pyropia in 2017 in China.
1.5 History of Pyropia utilization in China
The earliest record of Pyropia culture occurred during 960-1279 AD in Pingtan, Fujian Province, China. The ancient Chinese used lime to
remove attached algae and invertebrates from the rocks in autumn before releasing of ‘seeds’, which was named ‘Cai Tan’ farming method
(Fig. 6)[5].

Fig. 6 Ancient mariculture and harvesting of Pyropia in China.

1.6 ‘Caitan’ farming in China

The ancient “Caitan” method is still in use in some coastal areas in

a
South China (Fig. 7).

Fig. 7 Heavy P. haitanensis beds are distributed on the rocks in

South China (a) and the local farmer is harvesting them by hands (b).
1.7 The landmark for modern Pyropia industry

1. KM Drew’s finding in 1949: conchocelis has been determined as a life stage of Porphyra other than a separate alga species[6].

2. Ecological and developmental investigation on the life history made it possible to manipulate the life history in culture.

 Long preservation of free-living conchocelis for germplasm

 Large-scale culture of shell conchocelis for industrial producing (Fig. 8)

3. Establishment of raft-aquaculture. Two traditional culture methods for

Pyropia different from those for other seaweeds.

 Standing bamboo poles (Fig. 9)

 semi-floating rafts

Fig. 8 Shell conchocelis of Pyropia (Porphyra).

Fig. 9 Standing bamboo poles culture system for Pyropia (Porphyra).

2. Theory and technology of seedling culture

2.1 Life history of Pyropia

asexual
l m Most Pyropia sp. have sexual cycle with an
cycle
alternation between the gametophyte and the

sporophyte generation. Some have both sexual

k and asexual life cycles. A few species have

a only asexual life cycle (Fig. 10).

j b
c
sexual Fig.10 The life history of Pyropia (Porphyra).
cycle
d a: Spermatangium；b: Spermatium；c:
Carpogonium；d: Fertilized carpogonium；e:
i Zygotosporangium；f: Zygotospore；g:
Filamentous conchocelis；h:
e
Conchosporangial branch；i: Conchospore
formation and release；j: Conchospore；k:
h f Seedling from conchospore；l:

g Archeospore；m: Seedling from archeospore.

 2.1 Culture of Pyropia shell conchocelis

 Conchocelis penetrates into shell matrix, grows and develops into conchospore, which is the seed of laver for cultivation.
 All the other algae can’t penetrate and live in shell matrix.
 The contanimated algae are blocked outside the shell matrix and washed off periodically.
 Shell conchocelis is applied for industrial seedling culture (Fig. 11).

a b c

e f

Fig. 11 Culture of Pyropia (Porphyra) shell conchocelis. a-b, the shell conchocelis is growing; c: plat culture of shell conchocelis;
d: vertical culture of shell conchocelis; f: management of shell conchocelis culture: washing.
2.3 Disease occurs during Pyropia shell conchocelis culture

Disease occurs when the environment

conditions are not suitable, e.g., the
a temperature is very high, the light is too
high or two low and the water quality is
not so good. Yellow spot disease is the
most commonly occuring disease in
Pyropia shell conchocelis at high
temperature (Fig. 12).

b c

Fig. 12 Kinds of disease during shell conchocelis culture. a: yellow spot

disease; b: white spot disease; c: turtle cracked disease.

Treatment: immersed with several hours to several days of

 Freshwater
 KMnO4 (Fig. 13)
 ClO₂

Fig. 13 The shell conchocelis with yellow

spot disease is immersed with KMnO4.
2.4 Conchocelis development and conchospore germinating into sporeling

Three morphologically distinct stages occur during conchocelis culture: vegetative conchocelis, chonchosporangial branches and conchospore
formation and release (Fig. 14). Different culture conditions are required for different stages of conchocelis.

a: Vegetative conchocelis b: Conchosporangial branch c: Conchospore formation and release

d: Conchospore e: Conchospore germinating f: Two-cell sporeling g: Gametophyte blade

Fig. 14 Development of the shell conchocelis, conchospore releasing and developing.

2.5 Seeding conchospores onto culture nets

In autumn, mass formation and release of conchospores occurs. The released conchospores are collected and attached onto
culture nets and germinate into gametophytes. The collecting was done completely by hand (Fig. 15a). Nowadays, most of the
collecting process is done by machine (Fig. 15b, c). In the afternoon, the shell conchocelis is washed and water is renewed
(Fig. 15d).

a b

c d

Fig. 15 Chonchospores collecting process. a: seeding conchospores onto culture nets by hand; b, c: seeding
conchospores onto culture nets by machine; d: washing shell conchocelis and renewing water.
3. Therory and techology of Pyropia aquaculture
3.1 Therory of Pyropia aquaculture: based on the endurance to dehydration
 Most Pyropia species can endure desiccation. The blade becomes crisp sheet and recoveries after rehydration (Fig. 16).
 During laver cultivation, the competitive algae green algae (Ulva) and planktons can not endure extreme desiccation. The
periodically drying culture protocols are applied for laver cultivation so that the contaminated algae are killed by drying.

Fig. 16 Pyropia blades are naturally dried to crisp sheets on the coastal rocks during ebb tide.
3.2 'Green nets' and treatment

The nets full of green algae are called green nets (Fig. 17), usually occurred when the seawater temperature is high and the culture nets
failed to be effectively and thoroughly dehydrated. The dominant competive alga species is Ulva prolifra. On one hand, long period of
drying under blazing sun can effectively prevent the burst of green algae. On the other hand, the green algae can be killed by low
concentration acid and a period of freezing at around -20 ℃.

a b

Fig. 17 Pyropia cultures are full of green algae (a) and the green nets are dried on ground under blazing sun (b).
3.3 Pyropia is usually farmed in near-shore shallow water

Due to the highly dessication tolerance of Pyropia gametophyte blades and their easy suffering from the other competitive algae such
as green algae and diatoms, the culture systemes with periodically emerging from and submerging into seawater are used for Pyropia
cultivation. The semi-floating system (Fig. 18a) and the supporting bamboo pole system (Fig. 18b) are traditionally and the most
widely applied for Pyropia culture. The former system is applicable in shallow water with large tide range and the later system is
applicable in shallow water with depth < 20 m. Therefore, Pyropia is usually farmed in near-shore shallow water.

a b

Fig. 18 The semi-floating system (a) and the supporting bamboo pole system (b) for Pyropia culture.
3.4 Farming is extending to offshore deep seawater

The Pyropia culture areas have been expanding in recent years, however, the suitable coastal shallow areas are nearly fully
exploited for Pyropia farming. It is necessary to extend Pyropia culture to offshore and deeper seawater. The semi-floating
system and the supporting bamboo pole system are not applicable for deeper seawater cultivation. Culture systems have been
improved to support Pyropia culture in deeper seawater (Fig. 19).

Fig. 19 An improved culture system for Pyropia culture.

3.5 Farming systems for Pyropia culture
3.5.1 Semi-floating culture system

Fig. 20 The semi-floating culture system. a: the system is immerged at tide rising; b: the system is submerged at ebb tide.
 Characteristics and limitation of semi-floating culture system

When tide rises, the culture nets are floating and immerged in seawater. When tide ebbs, the culture nets are submerged and
dried in sun (Fig. 20).

The system includes bamboo supports, anchors and culture nets (Fig. 21).

The advantage of this culture system is that the culture nets are periodically dried along with the tide cycle and independent
of labor or machine. So this system is economical and manageable.

It has disadvantages: limited to shoal with large tidal range where there would be enough time for drying the culture nets.
Besides, it's limited to the areas where the low tide during spring tide should be daytime so that the nets can be effectively
dried. What's more, the time and period for drying can’t be manipulated, so the nets can only be passively dried.

Anchor

Culture nets
Support

Fig. 21 The basic structure of semi-floating culture system.

3.5.2 Supporting bamboo pole culture system (Fig. 22)

Fig. 22 The supporting bamboo pole culture system. a: the system is emerged at low tide; b: the system is hanged up and
submerged at ebb tide.
 Characteristics and limitation of supporting bamboo pole culture system

The structure is also simple with bamboo supports, anchors, culture nets and big and small floaters (Fig. 23). The advantage of this system is that the
nets are dried under necessity. The nets are hanged up and down on the supporting poles. It had disadvantages: it's limited to shallow water because
the length of the poles is limited; the nets are dried mostly by hand, so it's labor-consuming.

Bamboo
Small floater
Big floater

Anchor

Bamboo support

Culture nets

Small floater

Anchor

Fig. 23 The basic structure of the supporting bamboo pole culture system.
3.5.3 Full floating culture system (Fig. 24)

Advantages: Disadvantages:
• The nets can’t be submerged and dried.
• Applicable for deep water.
• Limited to large current and clean and clear water.

Fig. 24 The full floating culture system.

3.5.4 Improved full-floating culture method

This culture system can be turned over. The nets are growing under seawater (Fig. 25a, 26a). When it is turned over, the nets are
dried upon the big floaters (Fig. 25b, 26b). This technique has gained Chinese patent authorization (YSFRI).

a b

Fig. 25 Diagram of the improved full-floating culture system. a: emerged; b: submerged; c: the Chinese patent autorization.
a  Characteristics and limitations

Advantages：
• Applicable for deep water
• The nets are dried under necessity
• Easy management and labor saving
• Low cost for the facilities

b Disadvantages：
• The net is difficult to be turned over when laver
grows up and the quality of product would be
effected.

Fig. 26 Application of the improved full-floating culture system. a: when growing under
seawater; b: turning over to dry out.
3.6 Harvesting Pyropia products

When the Pyropia blades grow up to 15-20 cm in length, they are harvested by ships (Fig. 27). They are harvested for several times within a farming season.
Generally, they can be harvested every 10-20 days.

a b

Fig. 27 Harvesting Pyropia products. a: the harvesting ship; b: the big bamboo baskets used to hold the harvested Pyropia; c: the ship is harvesting Pyropia.
Limitation and problems of the current nori
industry in China
1. Limitation of germplasm
 Recruitment of new germplasms except for P. haitanensis and
P. yezoensis
2. Limitation of farming scale
 Already the largest farming scale
 Farmers and industries wishing to increase the scale
3. Farming technology
 Semi-floating and supporting bamboo pole limiting the culture
area to water depth < 15m
 Increasing farming scale by extending culture area from
shallow water to off-shore deep water through new farming
protocol
4. Seedling culture technology
 Shell conchocelis to be replaced by free-living conchocelis
 Hand work to be replaced by automated mechanization
 Issued nori cultivars in China
Cultivar Species Time Trait Protocol
Minfeng No1 P. haitanensis 2007 High output Selection breeding
Shenfu No1 P. haitanensis 2009 High output and high Mutation and selection
temperature-tolerance breeding
Shenfu No2 P. haitanensis 2014 Resistant to high Modern cell engineering
temperature, abundant technology, mutation
conchospore releasing, and selection breeding
high pigment content
Sutong No1 P. yezoenesis 2014 High output, Mutation, high irradiance
acclimating to high stress applied, and
irradiance selection breeding
Sutong No2 P. yezoenesis 2015 High output, Mutation and selection
acclimating to high breeding, modern cell
irradiance engineering technology
Germplasm collection
and preservation
 0.8 0.8 Stress response &
Female
0.7 0.7
Male acclimation mechanism
0.6 17 0.6
h
0.5 5d 0.5
Fv / Fm

0.4

Fv / Fm
0.4

0.3 0.3
0.2 0.2
0.1 0.1
0 0

Male
Female
Male

Male

Male

Male

Male
Female

Female

Female

Female

Female
-0.1
90.5% 68.3% 56.1% 45.9% 30.7% 23.5% 12.7%
LTLL LTML LTHL HTLL HTML HTHL AWC

WJ Wang et al. J Appl Phycol 2011, 2015

0.8

0.7 3h

0.6 3d
5d
0.5
Fv / Fm

7d
0.4
10 d
0.3

0.2

0.1

0.0

Male

Male
Male

Male

Male

Male

Male

Male

Male

Male
Female

Female

Female

Female

Female

Female

Female

Female

Female

Female
FW 7.8 15.6 22.6 29.6 36.6 43.6 50.6 57.6 64.6
Salinity
Identification of the genes regarding fatty acid
metabolism
Identification of the key enzymes regarding unsaturated
fatty acid metabolism
Identification of the key enzymes regarding stress signaling
Function categary Putative protein
Antioxidant ascorbate peroxidase
heme peroxidase
catalase
catalase/peroxidase
peroxidase
chloride peroxidase
cytochrome c peroxidase
cationic peroxidase 2
vanadium-dependent bromoperoxidase 1
Superoxide dismutase
Phospholipid hydroperoxide glutathione
peroxidase
Thioredoxin reductase
peroxiredoxin
1-cys peroxiredoxin
2-Cys peroxiredoxin
ferredoxin thioreductase
GSH-dependent dehydroascorbate reductase
Glutathione metabolism glutathione-s-transferase
glutathione peroxidase
glutathione reductase
glutamate--cysteine ligase
Glutathione gamma-glutamylcysteinyltransferase
3
Hydroxyacylglutathione hydrolase
S-formylglutathione hydrolase
Lactoylglutathione lyase
Glutathione synthetase
Flavonoid biosynthesis flavonol synthase
flavonoid 3'-monooxygenase
chalcone synthase
caffeoyl-CoA O-methyltransferase
anthocyanidin reductase
Other stress related ubiquitin and related proteins
ABC transporter
Heat shock proteins
Seedling culture of new germplasm
 Regulation

Within 15-20 d，
80% vegetative
conchocelis
developing into
conchosporangial
branches
 9000 A 25 0.6 A 0.6 B
a ab b
a
Pigment content (µg g-1 FW)
28 b a
b 0.5 0.5 a
31 a
6000 0.4 0.4

Fv / Fm
c
0.3 0.3

3000 b 0.2 0.2

a a a a a
a a b 0.1 0.1
a a a

0 0 0
APC PC PE Chl a Car 25 28 31 10 20 40 60
12000 B 10
Pigment content (µg g-1 FW)

Temperature (⁰C) Irradiance (µmol m-2 s-1)

a 20
9000 a 0.5 0.5 D
40 C c
b 60 0.48 0.48
b c a a

Fv / Fm
6000 0.46
b 0.46 b
b 0.44 b
0.44 a
3000 a a a b
a a b c c 0.42 0.42
a a b a a b c
0.4 0.4
0
APC PC PE Chl a Car 0.38 0.38
12000 C 8 L:16 D 25 30 35 40
Pigment content (µg g-1 FW)

a a ab 10 L:14 D
9000 Photoperiod Salinity
b 12 L:12 D
b
14 L:10 D
6000
24 L:0 D

3000 a a
aa
bbb abbaa
Ecological control on the conchocelis
a aa
aaaaa
development of new germplasm
0
APC PC PE Chl a Car

WJ Wang et al. J Appl Phycol, 2016

Farming of new germplasm
 The future of Bangiales (nori) industry
Construction of core
germplasm bank.
Diversifying germplasm
for farming, enabling
higher stress-resistance
Stress acclimation
mechanism：by innate
factors or by acquired
adaptations or by both.
Breeding of stress-
acclimating cultivars
Genome-wide high density
genetic linkage map
construction, functional
genes screening and
pathways research.
Molecular (assisted) breeding
Molecular mechanism of
conchocelis development.
Rapid and economic
seedling culture through
free-living conchocelis
Thank you for your
attention！