NOAA Technic I R

art

FS Clrcul

Proceedings of
the Sixth U.S.-Japan
Meeting on Aquaculture
Santa Barbara California
August 27 -28 1977
Carl J. Sindermann (Editor)
Und ( , e US Japan Coop (atl
I A soure s UJ R

P(m

In N tu(

rch 1

2

ERCE

2

PREFACE
The United States and Japanese counterpart panels on aquaculture were formed in 1969 under the UnitedStates-Japan Cooperative Program in Natural Resources (UJNR). The panels currently include speCIalists
drawn from the federal departments most concerned with aquaculture. Charged with exploring and developing bilateral cooperation, the panels have focused their efforts on exchanging information related toaquaculture which could be of benefit to both countries.
The UJNR was started by a proposal made during the Third Cabinet-Level Meeting of the Joint UnitedStates-Japan Committee on Trade and Economic Affairs in January 1964. In addition to aquaculture, current subjects in the program are desalination of seawater, toxic microorganisms, air pollution, energy, forage
crops, national park management, mycoplasmosis, wind and seismic effects, protein resources, forestry, andseveral joint panels and committees in marine resources research, development, and utilization.
Accomplishments include: Increased communications and cooperation among technical specialists;
exchanges of information, data, and research findings; annual meetings of the panels, a policy coordInative
body; administration staff meetings; exchanges of equipment, materials, and samples; several major
technical conferences; and beneficial effects on international relations.

William N. Shaw - United States
Masaru Fujlya - Japanji

I'
rqlO ITable l.--Species of food algae used to rear the larvae and juveniles of aquatic
animal s . Arrows indicate the direct utilization as food items to rearing species.

Species of algae
Chrysophyceae
Monas sp .

Method of utilization

Species reared

Crassostrea gigas
Anadara brought nii
Pte ria penguin
Spisula sachalinensis
Pinctada fucata
Haliotis gigantea
HaZiot'is d1~SCUS

Monochrysis lutheri

Anadara broughtnii
Atrina pectinata
Patinopecten yes soensis
Pinctada fucata
Pecten albicans
Chlamys nobilis
Fulvia nrutica
Spisula sachalinensis
Schizothaerus keenae

Bacillari ophyce ae
Chaetoceros calcitransAnadara broughtnii
Patinopecten yessoensis
Pe cten albicans
Chlamy s nobiZis
Fulvia nrutica
Mer etrix lamarckii

Chaetoceros simplex

Spisula sachalinensis
Anadara broughtnii
Schi zothaerus keenae
Halioti s gigantea
HaU otis di scus
Pseudocentrotus depreqsus

d.

l . --Co

io

d

sp .

, Zoo isp .

hom sp .

H io
NOlJi

~Za

sp .

H 'lioH 'lioCOOHilitz. hiaSf'>.

l.o.·t riwn

- - Gresp.

r---Table 1.--Continued.
Species of algae

Method of utilization

Chlorella sp.

Species reared-----Brachionus(marine)plicatilis -----l~~or

PagY'Us 17UJor
Opegnathu8 fas iotus- - Tigriopus japonicus----..

Acanthopagrus schlegelii
Kareius bicoloratu8
Limanda yrJkohaJrae
Sebastev inermis
Fugu rubripeo
Lateo labrax j aponi us
Parali hthys olivacus
Nep tunus trituherculatus- - - Green water - - Chlorella sp.

-

Daphnia pu lex ----l~~Marine fishes and shellfishes
Freshwater fishes(freshwater)

Table 2.-Composition of culture solution for Monochrysis luther;'
I,(XXl mJ
100mg
14mg
12.6mgIB.I mg
500mg
0.21'g100l'gII'

Seawater
NaNO,
Na,HPO'12H ,O
NaHCO,
EDTA·2Na
Clewat · 32·
VitaminB"
Vitanun B,
Biotin· Chemical composition: Fe, 3.B g; Mn, 7.7 g; Zn, 1.6 g; Cu, 0.07
g; Mo, 6.3 g; B, 24.7 g; Co, 0. 17 g; EDTA, a bit.

sheets of the collectors are hung in a tank tilled with eawaterand placed under suitable light; ub equently various species ofattached diatoms grow on the surface of the plate. The e collectors are transferred and hung vertically in the abalone rearing
tanks just before the planktonic stages of the larvae transform
into the creeping stage (Shibui 1971)
In the culture of red sea urchin, P. eudocenlrotus depressus,
phytoplankton species such as C simplex and S. costatum are
utilized for food of the larvae. The~e algae are cultured bymethods similar to those described for the culture of Pinctada
larvae (Hirano 1963).

INGREDIENTS OF
ARTIFICIAL COMBINED FOODS
Table 3.-Composition of culture solution for Nitzschia c/osrerium.
Filtered seawater
NaNO, (5 g/ lOOmJ)
NaHCO, (16.8 gil)
Na, SiO, (2 gi l)Na, HPO • . 12 H ,O (5 gil)PI. solution
Sterilized at 80 °C for 30 min

I,OOOmJ
2mJ
IOml
2ml
2m1

ImI

PI. solution
Na,EDTAFeCI , ·6H,D
ZnC!,
MnC!, ·4H,O
CoCl, ·6H,O
CUSO. ·5H,O
H, BO,
Distilled water

3.0g
O.24g
0.03 g0.27 g
0.8mg
DAmg

3.44g
l,OOOm!

Attempts have been made to utilize Chlorella pyrenoldosa
cultured in freshwater as an ingredient to increa e the comersion efficiency of artificial combined food for the creeping
larvae of abalone.
Recently, the production of Spirulina sp. has reachedcommercial scale. The mass culture of S. platensis and S. maxima
has been underway since 1974 by an industrial chemical company,
through a technical agreement with Institut Fran~ais de Petrole(Nakayama 1975). The company built a culture facility with 5,cxx:J
m' area at Miyakojima in the Ryukyu Islands. They expect toproduce 33 t (dry weight) of Spirulina annually. Because of highproduction costs, Spirulina has, in the past, been utilized only fordeepening the red body color of goldfish and fancy colored carpby taking advantage of its high carotenoid content (Honma et aI.
1974). In recent years, however, Spirulina has been utilized as an

I II H~It IH. ( I If J)

Recent Problem ofori (I) rp}lJ ra pp.
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• Kun; 2.- \" [j hoi 'l<'t'dong 01 ~on hTechniques of Culturerem Iron mental
tern

Maintenance of conchocelb cuiturl"l and collection uj
conchospores.-As the u e of artificial con~ho pore 011 tlon
has increased, it has become neee ar) to artilicrtlly maintain or
culture the conchocelis tage. Al first, the lage"a ultured 10
prefectural fisherie experimental staliom and l or other fl lillie
for the nori culturists. Today mo t cult uri t rai c theIr o\\n upplies of concho..:elis tages; therefore, the actual number of onchocelis cultures has greatly increa ed, bring10g about a problem
of insufficient supply of oyster hell ~uitable for u easubstrate. This has re ulted 10 the development of artifiLlai
substrates.
Recently, the free-living method of growing on..:hoceli tage
has been used by government research laboratone~ and pri\ atecompanies. This method facilitates genetic studie~ and breed 109
in the field, because a large number of conchocelis stage can beproduced from a single mother thallus. The dra\l, back to thismethod is contamination by diatoms and other seaweed pores.
Direct conchospore coUection has not yet been artificiallyachieved.
Seeding methods commonly used are "zubo" and "hanzubo." In the han-zubo system, tanks (I m x 20 m x 0.5 m
deep) made of heavy gage flexible vinyl are floated on the urface of the sea so that the sides are about 10 em above the water
surface. Each tank contains about 50 "net hibi" and about 500
oyster shells implanted with conchocelis stages. The zubo system
is similar, but is completely enclosed (Fig. 2).
Recently, a conchocelis net, the "zubo net" which has been
improved from the earlier zubo apparatus and made of smallmeshed double nets, is used for artificial seeding. At seeding, aconchocelis net with about 300 sheUs, which have been previously arranged within it, is set on a framelike simple raft; usually
about 50 nori nets to be seeded are spread and staked on theraft. Finally, the raft is covered with a vinyl bag, but inside calm
bays the raft is often floated without a cover. The raft is then left
about 3 d to achieve good implantation on the nori nets.

reMaintenann' 01 ju ~rOlI to

used Unl\er 11) In J p n lod
ru ture .
I) The" 01C! hi ." (or ft ed melh ) I U
In h 10 andaim \l,oler • e g , \l,11h1O a ba) \ here pol are em~dedin the bOllom.
2) fh "betan g hi" ( no lin) 1 U
and 10 the open ea here \ nou buo) nI object pro\,notation and an ho m nl In the po IlIOn nd hape 0the ) tem.
There i a third mt"thod \\ hI h I 1Olennrotalt" between tht"
abovt" t\l,O mel hod . It IS called "han rudo hI I" ( half f} tin
method) and i u ed "here there i alar e IIdal tluctu uon. Thi
method IS u cd 10 avoid exce I\e ur),ng dunng 10\1, tiut" penod •but keeps the Porph)'ra at tht" \\att"r urfaee for rna lmum light.
The mo t important diflcrenLe bCI\\een kott"l hikl andbetanaga hi, a far as Porphl'ra spp. arc concerned, i the prO\1sion for drying durmg cultivation. Thalli gro\\ n under nondr)ing conditions art" more u ceptible to di ea e. and Ihe nori produced without a df);ng penod are nOI as high in quality a thenon grown with a dr);ng pt"riod. Ho" t"\ er, to date it has no
been scientifically substantiated that dr)ing is the dlrt"Ct cau eo
the increase in qualJt>.
In the past it was not possible to allo\\ a controlled dr)ing
period using the floating raft melhod. The juveniles \l,ere
cultured with the fixed method mshore and then tran planted 10
a floating raft offshore 15-30 d before harve t. Recently M. Inayoshi, a nori culturist, improved the betanagashi method b)
adding a structure which facilitates raising the nets out of tht"

8

Figure 4.-Frozen non nels stored in - 20"(; freeur. on nets are packed in a vinyl bag afler semJdrying and thenlhe bags are sealed, since Porphyra should be protected from fresh aJr to mJnimize their respinllion.

Figure S.-Non harvesting with a rotary blade, suction-type harvester.

against diatom propagation, such as Ge02, were discovered(Kirita 1970; Kida and Enomoto 1975). The entire free-living
technique is still experimental and requires improvement before
transfer to mass culture.

grow healthy young thalli of Porphyra. Moreover, the sheets
made from the thalli which have undergone the drying process
are of better quality than the ones grown without it. However,the drying results in a lower growth rate of the thalli. Considering these facts, the improved floating raft should be
employed extensively throughout Japan, and physiologicalstudies on the effect of drying the thalli should be continued.

Improvement of the Maintenance Technique from
Juvenile to Adult Thalli
It is well known by the cultivator that drying the growing Porphyra is an effective way to remove contaminating seaweed suchas Enteromorpha spp., Ulva spp., and diatoms, and also to

Development of Effective Species and Strains
The quality of the two most popular species and their newtotial dangers, the priority of water quality must be recognized by
all concerned.

CONCLUSIONS
Although nori cultivation is almost standardized in Japan,
there still remain a few constraints. Nori is marketed a\ raw
sheet nori or processed nori . At present, the demand for raw
sheet nori (which is premium nori) is greater than the supply . On
the other hand , the supply of proce sed nori is greater than the
demand.
In view of the present situation, the future success of the noriculture industry can be best achieved by encouraging increaed
consumption . This can be done by maintaining the highest
quality in the product and lowenng production co ts . ew
technological Improvements could influence and in ure highquality. Production costs can be lowered by changing or improving existing practices through cooperative efforts similar to
those used in agriculture : I) cooperative cottage industr), 2)
modified cooperative unions, and 3) full -scale cooperative un ion
management. An economic evaluation of the Industry would
also be desirable.

States-Japan Cooperative Program in alural Re!.ource5), andRobert D. Wildman and John H. (,lude, OAA, lor their kind·ness; Tim Ahern, University of Wa hington, for hi kind help
and asmtancc with f!nglish: and Shigekat u Sato and Hi hi
Kanno, Tohoku Regional h herie Research I.aboratory, fortheir support.

LITERA" URf, CITED"IDA . W .. anu T. [O M ol0197
O n the protectl n from I contaminatiOn In Ihe culture 01 conju epores 01 'tofonuSlromo lin Jm J Pr
nn r"ICCI Jpn
1M
Soc. Hi> J'
KIRITA ....1.
of I'orph rro'970. I ff«1 of), on the pO'Oth 01 onch mu p
(In Jpn I Bull Jpn f'hy I Soc I )1 I , 170
197 . . ....\
cullure 01 con h cliJ phue In free livin form and dlrca
Ann i CClJm
collectm Icx.hmquo 01 \.On h pores
IIn Jpn I
Sci l uh Soc. 47 P

I',

\I URA , ,\1'17U
peoo anu Jtnuns 01 Porp TO
lor nonlIn
Jpn I / " Saburo Ucda, cdllcd by.
ataYulllClu no non en yu (Ournon cull Ie Jtudy), p S 2~e yolume In I ,allona!
I edCl' lion 01
n and Clams r Mm C pcnllvO. To yo
1973
Reeent nor! production Incrcasm In the rnmncr of ~nc pr~plon, and Ihe sclf-conu dm.1OO appeared In the c(In
Jpn.1 l:colo81cal lecture ('ieUm
KouJ. Month!) ,cpon ' 17. p i )J..: YOt'U hupp:m, To yorl\hm

ACKNOWLEDGMENT
The author exp resses deep appreciation to William . Shav, .
chairman of the U. S. Aquaculture Panel of UJNR (U nited

UI!DA.
1911.

:-,,,,, ~cm for nor! ulture (In Jp:o I
1I0nal Fcdcra JOn of\arm fMm <. per • ,To •• p I

n andu ____________________________________

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nt tatus of Brow nInl

I~ I HOIII
I nd ,/.7

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nil I llllinori l•.,ern Japanese coast of Hon~hu. The annual harvest from natural
beds is estimated at several thousand metric tons (wet weight). Its
life cycle, ecological characteristics, and cultivation techniques
were studied by Migita and Yotsui (1972), Yotsui and Migita(1974), and Yotsui (1975a, b, 1976) (Fig. 5).
Cladosiphon okamurans is found chiefly on the skeletons ofdead Madreporaria and seaweeds, sometimes on pebbles, cans,
sheIls of dead shellfish, and other hard substrates. Its natural
distribution is limited to the Ryukyu Islands. The annual harvest
from natural beds is estimated at a few thousand metric tons (wetweight). Its life cycle, ecological characteristics, and cultivation
techniques were studied by Shinmura and Yamanaka (l974a,b) and Shinmura (1974, 1975, 1976) (Fig. 6).
The cultivation of these two species involves similar methods,
except for the water temperature necessary for spore release,germination, and growth. A characteristic difference between th
two groups of cultured algae is that cultivation procedures includ
the entire life cycle for Porphyra, Undaria, and Laminaria, anonly part of the cycle for Nemacystus and Cladosiphon. In
autumn, the gross sporophytes bearing plurilocular sporangia collected from the field (and also small sporophytes that have been
cultured in containers since spring) are placed in tanks under illumination of 2,000 lux with gentle aeration. After a large numbe~of spores have been released into the seawater, culture nets madeof synthetic fibers are put inside the tank to collect the zoospores
After the collection of zoospores, the nets are left in the same tank]for 1-2 d and then transferred to sea, where they are spread inwater of about 0.3 m deep for Nemacystus and of about 1 m deen
for Cladosiphon. The culture nets are supported in the desired
water layers by buoys and held by anchors, as shown in Figure 7
A few months after the collection of zoospores, Nemacystus growto 10-30 cm long and Cladosiphon to 20-40 cm long unde~

Kt

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a

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a

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0.

figure 3.- AnnuaJ variation in production in Laminaria cultivation.

Under these circumstances, managers of Undaria andLaminaria culture operations are expected to increase production
per cultivating unit and improve the quality of products. To attainthese goals, it is necessary 1) to improve techniques to prevent
infections and competition from harmful organisms, 2) to haveefficient field fertilization, 3) to breed excellent strains, and 4) toculture healthy seedlings.

Nemacystus and Cladosiphon
Nemacystus dicipiens grows on sargassaceous algae, especiallyon branches of Sargassum patens, and is distributed on thesouthern Japanese coast, the west coast of Kyushu, and the north-sept.
Two-yearfarming

Oct.

Nov. Dec.Jan.

Mar.May Jul.Sept.

Artificial
seedling

~

.c
.j.J

Jul. sept.

I - - - - . t Second year

cultureof culture)( Flo~tingdev~ce

a
o

".-<
t1>

Nov.Jan.Mar. May

Reject of)( fouling organism
Transplantation
farming~oRepairing of )( holdfast(Indoor hardening culture)z

Transplant. ofthinned Laminaria
Transplant. of
running 2-year
Laminaria

Forcedfarming(Indoor forced culture)(Fertilization)

Hastened Laminaria 1 - - - - - - 'cultureFloating of culture)(Thinning)( device
Repa~ring( holdfast
Mi d-May

Jun.

Mid-May

Oct.

";

a
o

One-year
Artificial seedling Indoor culture
forced farming L -_ _ _a-_ _ _-.-J ' - - - - - - - - '~(Indoor forced culture)

Nov.

Of)

Nov. May!

DryingReject of)( foulinq organism

.c
.j.J
:>

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Figure 4.-Laminaria cultivation process flow diagram.

14

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Marketsmall mesh net, washed with seawater, and suspended in clear
seawater. The embryo suspension is then sprinkled onto the seedling twine which is wound evenly on a square plastic frame placedin a shallow tank. The material used for seedling twine is mostly
synthetic fiberspun yarn, about 2-3 mm in diameter. Two or threedays later, the frames are removed and hung vertically in a deepertank . When the buds grow to 5-10 mm long, the frames are tak~from the tank and suspended in the sea from a raft. Here thframes remain until the buds develop into healthy young th .
about 10 cm long (Tomiyama 1974; Yoshida and Nishikawa 1975;
Yamaguchi et al. 1977; Toma et al. 1977).
These propagation procedures are often unsuccessful owing topredators. As shown in Table 2, various predators on macro algae
are found in Japan. It is not desirable to kill all species 0
predators prior to the establishment of seaweed beds; since some
useful species are present and interactions are complex, it inecessary to keep the biological balance between aquatic faunaand flora in the area. To achieve this, three methods have been
tried: 1) To capture or kill unused herbivores, 2) to supply usefulherbivores with additional food by cultivating a large amount oti
algae at or near the sea surface and then to harvest these herbivores after they reach a suitable size (Kikuchi et al. 1975), and 3)
to use nets to cover the newly established substrates to whichseedlings are attached, until the seaweeds have grown to a certain
size, as shown in Figure 8 (yamaguchi et al., 1977).
For the further development of brown algae propagation in the
sublittoral zone, it is necessary to: 1) further clarify thebryos. Sometimes they are shaken in the seawater to induce therelease of embryos. The collected embryos are ftItered with a

Table 1.-Spedes of brown algae used for afforestation.
Methods of recruitment
of seedlings
Transplanting
culturedseedlings

SettingmotherfrondsSpecies nameLarninariaceae
Laminaria japonica
Eisenia bicyclis
Eisenia arborea
Ecklonia cava
Undaria pinnalijida
Cystoseiraceae
Cyslophyllum sisymbrioides
Sargassaceae
Sargassum piluriferum
Sargassum patens
Sargassum pinnatijidum
Sargassum horneri
Sargassum serratifolium
Sargassum lorlile
Sargassum fulvellum
Sargassum ringgoldianum
Sargassum Ihunbergii
Sargassum assimile
Sargassum si/iquosum

o

o

o
o

o
o
o

o

o
o
o

o
o
o

o
o
o
o
o
o

o
o

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ill

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414 14 I 4I 4I •t

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Figure 8.-Net for protecting young buds, for example, from bannful grazers (Yamaguchi et aI. 1977). A. Square net (mesh size 2em). B. Frame rope for fence net (2.4 em diarn .). C. Anchor rope (1.8 em diarn.) . D. Concrete anchor. E. Fence net (mesh size
1.2-1.5 em). F. Underwater zipper. G. Lead chain sinker.

16

he

se of Phytoplan ton for qu cultur
a tatu R port

I HOIH ( 110Figure 2.-Phytoplankton tanks at a commercial shellfIsh hatchery on Long Island, N.Y.

Phytoplankton production plays a key role in the success or
failure of a shellfish hatchery. It is one of the most expensive
items associated with a hatchery and takes a large amount of

space. Recognizing these factors, studies are underway in several
laboratories to replace phytoplankton with artificial diets.
Studies are also underway to utilize phytoplankton, not only
for the hatchery aspects of bivalve culture, but also for growing
mollusks to marketable size. The University of Delaware is
growing large volumes of algae, principally Thalassiosira
pseudonana, as part of their attempt to develop a controlled environment for shellfish mariculture. Initially, the algae weregrown indoors, utilizing artificial lighting (Fig. 4). The highcosts for artificial lights needed to produce large volumes ofalgae caused the researchers to switch to an outdoor greenhousesystem (Fig. 5). In recent studies (pruder et al. 1978), cell counts
in 9,000 liter containers have been increased from less than 2 X
lO'/ ml to over 10 X lO'/ ml [a 5-fold increase) by bubbling N,
and CO, gas mixture. The addition of gases controls the pH anddissolved oxygen levels which in turn favor algal growth. The
main difficulty is finding suitable algal species that can withstand high temperatures and also promote shellfish growth. In
earlier experiments with two species of algae, T. pseudonana
and I. galbana, oysters have been grown to market size in 9 mo.
Unfortunately, the latter algae cannot be grown undergreenhouse temperatures. Recently, another species ofIsochrysis from Tahiti has been grown successfully in the controlled environment and appears to be an excellent food forbivalve mollusks.
In St. Croix, Virgin Islands, a unique culture method is being
tested, using nutrient-enriched water pumped from 800 m
depths to grow phytoplankton. Deep water is pumped into two
50,000 liter pools (Fig. 6) that have been inoculated with
Chaetoceros curvisetus (Fig. 7). The water is then pumped continuously to raceways containing the clam Tapes japonica. Thissystem has produced excellent growth and a high quality product. The economics of the system, plus final testing of raceway
design, are now being examined.

• I~un'Ph\ lupl:Jn"'lon h~tn2 poured Inln ",helltl... h Ian ae ~ro"'-outlan"-,.

In Woods Hole, Mass., studies are underway using the effluent from secondary sewage treatment, which is mixed withseawater to grow a single-celled marine alga in 120,000 liter
ponds (Fig. 8). The seawater containing the bloom of

I'"

4

flKure 6.-Ph,tophlnklon POO~. 'I. (m \. \ 1'1[10 I I~adnece ry amount of phytoplankton: FeruhLJng the Ian.al tan .
to obtain the necessary bloom, or grov.,n the phytoplan ton
eparate1y and addll1g It to the culture tan • The prinCIpal
pecle cultured i kelelonema P. although other pa1 • u h
as TelraselmlS p and Thaloss/Os/ra p., ha\e al 0 b n u cd u~ce fully (Yang I). Recently, a pmate firm has been ulturing and freeze-drymg Skelelonema. Thi product ha been u ed
ucce~ fully in ultunng hrimp Ian ae III e as. :-'Io<:k (1974)
has ue e full} concentrated kelelonema b centrifuging,refngerating or freezing, and then uSing It at a later dale to feed
shrimp larvae.
Con Iderable intere t hde\elope
in the u e ofph}toplankton in culturing e\eral peci of Hacrobrachlum
larvae, particularly \1. ro enbergi/o FUJlmura (1966 tre ed thevalue of groy, ing AfacrobrachlUm Ian. ae in "green water." Heclaimed that algal supplement increased Ian aI un;, aI andshortened the lime to stockabk po tlan.ae. tudic<; b) ~1addo\
and :V1anzi () 976) concluded that algal upplements incre;c,ed
larval survi>a1 and the production of po llan ae \, hile decreasing
the length of time required lor metarnorpho i . In the above experiments, the 1an.·a1 tanks were inoculated with algae to produce cell counts ranging from 400,
to 1,3Iml.
However, a commercial company in Puerto Rico successfully
rears Macrobrachium lanae \\ithout algal supplement.
Therefore, the question of the need for algae for rearing
freshwater prav.ll lan·ae still remains unre olved.
Algae pia} another interesting role in the culturing ofMacrobrachium in ponds. The Puerto Rican company mentioned above has run into serious problems with aquatic weeds. In
an attempt to control the growth of weeds, fertilizers have been
added to the ponds to create an algal bloom. The bloom in turnacts as a shadow which cuts the sun's rays; thus, the aquatic
weeds cannot grow. The greatest problem encountered with thismethod is maintaining a stable bloom.

CRUSTACEAN CULTURE
Phytoplankton plays an important role in the nutrition of
shrimp larvae. There are two methods used to obtain the

FINFISH CULTURE
Experiments have been carried out in Alaska to test thefeasibility of using artifiCIal upwelling to enhance primary

Figure 7.-Cylinde\1i of phytoplankton used to inoculate pools, SI. Croix,
Virgin Islands.

22the 8,ant pm"n. \fa rohro(nlum m n"agll«(luncll. r"c1flh ScMlon. Honolulu. lIawall. 11'1

(Roberts et al. 1976), thread herring, Opisthonema oK/mum(Richards and Palko 1969), scaled sardine, HarenRu/a pensaco/ae (Houde and Palko 1970), and Florida pompano,Trachiontus carolinus (Hoff et al. 1978), thi procedure is used.
Roberts et al. (1976) also found that Dunallella or Nannoch/ons
algae maintained water quality when culturing C. me/ana.

"rIC

Indo-I'I W1ai'l"'/V.!'ol7. P

HOI I, (. H. J . 1011 I IN. I I IV\KI • nd K II I ()n197H . Spa"nmg. oocyte development and larwc rearml of lhe /lomb
pomp no (!rurn",,,(,, ('uro/mw)/" J '"
voult. 1r (edllor).
I'rOCtt(hnof Ihe '",h Ali j I leel n "'orld Marlcuilure '><xlClY.
J1 27'12'17.
!lOll!)1 • I !). and II I 1',\1 I\IJ
1'170 I borutory re-dTin of Ihe ciupe1d (j h Ifurl!nllu/a pmJDroia fromferhilled~Iar 11101 (Berl) ~ ~3~"'ADI)( lX. "I II . nd J J 1 1.1
Ihc effrc!
f III

SUMMARY
I have briefly outlined examples of the key roles thatphytoplankton plays in aquaculture. It is, in some ca es, one of
the major costs in an aquaculture facility. For example, in ashellfish hatchery in Maryland, over 200 liters of algae are useddaily to culture oyster larvae. The manager stated that thl
represented 40010 of his total productIOn co ts (Kranll') . To
reduce these costs, programs like the one in Delaware are u ing
sunlight and gases to increase cell counts. In other cases, hJ..e the
one now being carried out at the Virginia Institute of 1arine
Science, artificial diets are being developed to replace the algae.
Yet, it is obvious that phytoplankton will continue to be important in many aquaculture programs A substitute is till III the
distant future.

R BI RT<;. D E.

• and

R

LITERATURE CITEDt"

ALLE ,G. H
1976. Rearing Pacific salmon in ,ah"ater pond lertlhzed 'Alth dom II
wastewater October 1975-August 1976. Humboldt Statc L~v
urant
Program Rep. HSU-SG-Il. 99 p.
CASTAGNA, M
1975. Culture of the bay scaiJop, Argopec(en IrradwfI5. In \ Irglma :>'Iar
Fish. Rev 37(1): 19-24.
FUJIMURA. L
1966. Notes on the development of a pracucal ma.>S cullunng techruque for

R~THf:R. J

19n

HPreiumnary rmll
tern
In F
4. P
112

lar

IX

cr. I

(onfcren,~162(, V. T1975
manual for br Nan culture of prnaeId hnmp 10 Ihe posIlarval lag~
OJ\ \Lmu '>ell Gram. fech Bull 31.
p'George E. Krantz. University of Maryland Hom Point Environmental
Laboratory. P.O. Box 775. Cambridge. MD 21613. peTS. commun. January 1979.

24

R

J HOIH ( 110and onomi .. ~ ording to Huguenin (1976). !.he major problems and uncertrunties regardmg mten ne seaweed cultivation
are tho e of ale, hydraulic. re idence time. nutrient upplies,
monltormg and control of the facilit). control of epiphyte.
sea e· ph iol08Y, and species election. Basically an interdependent . no~ledge of seaweed biologj, engineering.
economIc. and operating poliCIes \l.iU be required for any ucful seaweed culturing ystem.
Hanesung Irom 10 ely managed natural stocks IS becoming
in rea mglj e pen Ive (e.g., with Chondrus crispus), and it may
not provide uffident material for the projected needs of commer lal marine plant indu tries. Hence. mariculture techniques
are being developed in Canada Neish (1976a. b. 1979) gave acomprehcn ive review of the CanadIan seaweed mariculture indu tne , which re in an early tage of development. Seven
trate 'e are being examined in order to enhance control overraw rna enaJ: I) ~1anagement of natural stocks, 2) addition ofub trata and habitat improvement, 3) live storage of han'ested
plant, 4) vegetative ropagation of wild tocks. 5) vegetative
propagation 01 elected clones. 6) grow-out of hatchery seed, 7)
gro .... -out 0 hatchery eed from selected seed.
T e Implest form of plant mariculture In Canada consists ofprovldmg ultable gro .... ing conditions (i .e , substrata) for commerciallj de irable pecles . Habitats may also be "improved"by the remo\al 01 undesirable specie and by fertilization ofnatural bed To date, neither of these strategIes is used extenively 10 Canadian mariculture programs. Live storage ofha e ted plants has been practiced for a number of years in thedul c, Pa/mana pa/mara; rock weed , Ascophyl/um and Fucus;
and In h mos indust ries . This may vary from simply holdIng
hane ted plant In bags in the sea for a few day until suitable
dry1n .... eather occurs to a complex form of holding hving
m tena) in large tan ks with agitation by compre sed rur. In suchtank, dul e can be kept alive and growing for 4-6 mo . The\ etau\e propagation of su~pended culture of eaweeds was
fir t mill ted in anada with C. crispu.s ( eish and Fo\ 1971;
CI h anha klock 1971; Shacklock, Robson. For yth and
I h I 73, ha klo k, Rob on. and impson 1973) at the Sand\
field tat ion of the atlOnal Research Council of\eral om panic (~1arinc Colloid, Inc . and Genu
Produ tnada, Ltd.) tla\ e re ently initiated .:ultivation pro,m of In h mba cd on this technology. as well as the
I II n f f t-gro .... IO' traim of C crispus. to increase yields\era! e. \\eed C:U1not be propagatednd r \\Ih r t
et u\el , r \\-out technique mu t be initiated. particularly
Ith nnu I pee!
r r<=CIC with an annual frond and a perennih Idf
of no mten I\e manne plant mari~ulture
nbed (. I h 1976a, 19 9), induding thekno\\ ledge of algal phy iology, and correlation and utilizationof exi ting knowledge .
Den ity and tank agitation are interacting parameters
as ociated \\ith propagation of detached seaweeds and have profound effects on yield, crop cleariliness, production cost, andother economically critical variables (Neish and Knutson 1979).
According to the same investigators, the maximum sustained,ields per unit of pond area occur at densities of 1.0-2.5 kg oflive algae m'. At lower den ities. fast growth rates were observed
but area yields diminished and epiphytization became a p~oblem .
Agitation by mechanical . pneumatic, or hydraulic means
reduced epiphytization by mixing the crop and by physicallydisrupting epiphytes. Agitation was also important from thestardpoint of mixing nutrients, exposing all fronds to light, andcontributing to the general homogeneity o f culture conditions.
An optimization of pond agitation, which requires energy, anddensity-yield relationships will obviously determine the capitalintensity of a culture system.
A general review of the principles and perspectives of seaweed
cultivation in enclosed systems has also been summarized (Neish
1979). The main features of closed systems are discussed in relation to several research and development projects in the UnitedStates, Canada, and elsewhere; i.e., instances where seaweeds
are being employed in polyculture systems, as well as for foodand fodder, as feedstock for chemical industries, as effectors oftertiary effluent treatment, as recovery vectors for valuable
compounds, and for removal of deleterious compounds ¥eenumerated. Overall, Neish characterized closed-systemseaweed cultivatIon as a hIgh-investment , high-technology
business requiring a firm foundation of scientifically verified
facts. An opportunity brief descri bing many of the technical,
economic, and biological factors associated with closed-cycle
aquaculture was recently summarized (Anonymous 1977d).
The seaweed cultivation program at the University of California, Santa Barbara, was recently summarized by Neushul et al .
(1976), Neushul et al. (1977), and C harters and Neushul (1979) .
They emphasized that many of the pro blems of creating asuitable environment for the growth and reproduction of large
seaweeds were yet to be solved. Successful experimental cultivation depends on having large volu mes of seawater, thetemperature and chemistry of which must be held withinprescribed limits. Contaminants and epi phytes must also be controlled (Polne et al. 1980). The requirement of water motion isparticularl, Important for many seaweeds (Charters et al . 1972,1973). The anta Barbara cultivation system consists of an outdoor greenhouse wllh aquaria, which is combined with an indoor
cultivatIOn room. having controlled light intensity andphotoperiod . The greenhouse was de~igned to permit the su tained growth 01 large kelps. several of whic h have been grown for0\ cr a )-car in a contInuously healthy state. Other plants have beengrow n from gl'rmlings. transplanted to the ocean, and periodically returned to the greenhouse for measurement and stud y.
Sc\cral othcr dcscTlptions of experimen tal culture fa cilitie.... ere given at the international symposium "The C ultivatio n o f
1anne Organi m and It Importance to MarIne Biology"
Kinne .lOd Bulnhelm 1970) f'or example, South (1970) described,U)e peTimenl.li culture lacllIty lor the grO\q h o f kelps,hI h on i ted of a t nk y tern supplied .... Ith fi ltered , freen w1ng cawater and artdi~ialillumination. He was a ble to o btam P:)[O ph)1c of ,.l/urta (,H ulentu 30 em long in 6 mo. startIng
por. I he paper b) Kornmann (1970). Jo nes andfrom z[) nt (I ~ ), ImonctlJ ct I. (1<) 0), Hnd Smith and .l o nes (1970)
fr m thme rnpo 111m h uld ul 0 be onsu!ted as they giveChondrus crispus (Irish Moss)

uneconomical, i.e., their cultivation requires too much labor.
An epochal improvement in techniques will be required to maketheir cultivation practical. The net cultivation studies of Iridaea
cordata by Mumford (1977a, b) should be noted, as he is attempting to obtain the data necessary for an economic analysis.
A variety of factors are important in determining the enhanced
utilization and development of marine plant resources (Scagel1961): I) Accurate estimates of quantities of economically important species; 2) determinations of the distribution ofunderutilized species; 3) investigations of appropriate methods
for harvesting; 4) growth studies of commercially important
seaweeds; 5) observations of seasonal variations in stocks andthe recovery of beds after harvesting by different methods; 6)
evaluations of chemical composition of seaweeds at different
seasons, habitats, depths, ages, and environmental conditions;7) correlating management measures with life history details;
and 8) introducing an educational program to acquaint the
public with these resources and their wide range of uses.
The status and potentials of marine agronomy have beenreviewed by Doty (1 979a, b, c, In press a, b); much of this information is relevant to the present account on seaweed
mariculture. He emphasized that marine agronomy programs
should be seriously considered by any nation bordering the sea
in view of man's needs and the fact that the territorial waters areas productive or more so than the adjacent land . Accordingly, aresource utilization plan must be initiated in order that the longrange gains will offset the losses. He recommended a broad
range of understanding of seaweeds, including an identificationof habitats available for their growth and information on theirnatural products. Such a broad base of information will provide
a vehicle for resource optimization based upon scientific algal
agronomics.
The need for a national mariculture research and development program in the United States has been outlined by Hanson(1974a, b), including four major areas of research and development: 1) Environmental conservation and predictive modeling;2) biological research on the selection of organisms, hatchery requirements, primary productivity, nutrition, husbandry anddisease, competition, and predation; 3) technological researchand development dealing with containment and attraction, concentration and harvesting, feeding technology, and artificalupwelling; and 4) socioeconomic research dealing with legal andpolitical questions. Suggestions concerning the appropriate rolesof government, industry, and the academic community withrespect to this proposed national program (United States) werealso discussed (Hanson 1974a, b; Anonymous 1978a).

Both the biology and ecology of Irish moss must be
understood if it is to be maintained as a natural resource (Taylor
1972a). Such information is equally fundamental fo r the successful mariculture of C. crispus. A synthesis of information
regarding Irish moss was recently summarized in a symposium
dealing with C. crispus (Harvey and McLachlan 1973). Subsequently a variety of investigators in the United States andCanada have further expanded our knowledge of the growth(Taylor 1972a, b; Prince and Kingsbury 1973a, b, c; Taylor et al.
1974, 1975; Mathieson and Burns 1975; Hanic 1977; Chen andTaylor 1980a, b), reproduction (prince and Kingsbury 1973a, b;
Cheney and Mathieson 1978b; Craigie and Pringle 1978;
Mathieson In press; Tveter-Gallagher et al. 1980), harvesting effects (Patell 1972; Taylor 1972a, b; Scaratt 1973; Taylor et al.
1974, 1975; Mathieson and Burns 1975; Pringle and Semple 1976,
1978; Cheney and Mathieson 1978b; Pringle 1979; Pringle et al.
1979; Pringle and Jones 1980), ecology (Bums and Mathieson
1972a; Kent and Buggeln 1972; Prince and Kingsbury 1973a, b, c;
Mathieson and Bums 1975; Taylor 1975b, c; Craigie and Pringle
1978; Lubchenco and Menge 1978; Pringle and Jones 1980), life
history-sporeling development (Burns and Mathieson 1972a;
Chen and McLachlan 1972; Ring and Vadas 1972; Tveter andMathieson 1974, 1976a, b; Chen and Taylor 1976; TveterGallagher and Mathieson 1980), outplanting techniques (Hanic
and Pringle 1978a, b), resource management (McDougall 1972,
1974; Pringle 1976, 1979; Pringle and Sharp 1980), carrageenan

MARICULTURE AND ECOLOGY
OF SEVERAL SEA WEEDS
This section summarizes current mariculture activities with 12
economically important seaweeds, including 10 red algae (Chondrus crispus, Eucheuma, Gracilaria, Hypnea, ·Iridaea, Gigartina, Gelidium, Pterocladia, Palmaria palmata, and Porphyra)and 2 brown algae (Laminaria and Macrocystis). A detailed
compilation of information on their cultivation, as well as
biology, ecology, and resource management, is given for these
economically important seaweeds in North America, the UnitedStates' possessions, the Trust Territories of the Pacific Islands,
and the Philippines.

A dnlg nlck used for dredging Irish moss in the Canadian Maritimes .
P hOlognlph by J . Pringle.

28feets of aeration, addition of nutrients, and variation ofphotoperiod and light intensity were also evaluated. In
evaluating the application of their work to commercial cultivation of Irish moss, Neish and Fox stated that it would not bd
profitable as a greenhouse crop, nor in a system employing sup]plemental artificial illumination. However, cultures in outsid
impoundments might be profitable from May to September atheir latitudes. Adequate nutrients (particularly nitrogensources), controlled access to seawater, temperature> lO oC, andthe availability of superior strains would be essential .

Recent stucties by A. eish and associates at the National
Research Council of Canada in Halifax demonstrated the potentials of Irish moss cultivation in greenhouse tanks flushed withseawater (Neish 1968; Neish and Fox 1971; Neish and Shacklock
1971; Anonymous 1977b; Brochu 1977; Neish et al. 1977;
Simpson et al. 1978; Simpson and Shacklock 1979; Simpson etal. 1979). For example, Neish and Fox (1971) showed that
detached macroscopic C. crispus plants grew in a greenhousewith flowing seawater and a temperature range of 5 °-15 °C, withgrowth being most rapid at the warmest temperatures. The ef-

Individual tanks employed for "seed-stock" selection of fast-growing strains of Irish moss at the greenhousecultivation facility of lhe Marine Colloid, Inc., Metagan, Nova Scotia, Canada. Photograph by D. Cheney .

.... , ",I pl .. nl

IIf

Ih

I ~ lUItU~ Inlln of (hllndrul crilpu\ IImh mo",) gro"lng in lank., at the Jack,on
nnr I ~b"",lon. :-.." Hamp,hor . Pholograph b) O. (hene) •I . IU.10

Seaweed culture ponds fo r the Irish moss cultivation program of the Marine CoUoid, Inc., Metagan, Nova
Scotia, Canada. Photograph by D. Cheney.

ject has attempted to produce strains of Ch ondrus with high carrageenan yields and/or growth rates, as well as to select featutes
potentially advantageous to cultivation, such as the ability toreproduce vegetatively by cloning. Chondrus populations froma variety of habitats (intertidal, subtidal, open coastal, andestuarine) and geographical locations have been compared forgenetic variability and differentiation, using starch-gel electrophoretic techniques (Cheney and Babbel 1975 , 1978; Cheney
and Mathieson 1977, 1978a). Such information is being used to
select diverse parental stocks for hybridization and the production of progeny which hopefully may show hybrid vigor . Electrophoretic screening (Adams and Allard 1977) is also being
used to recognize polyploid plants that may exist in nature.
Attempts to induce polyploidy in C. crispus are also being
carried out by Cheney and Mathieson using colchicine, which
inhibits the production of the spindle apparatus during cell division, causing a doubling of chromosome numbers. It is hopedthat a 2n gametophyte can be produced , which in turn will becrossed with a normal (1n) gametophyte, producing a 3n plant.
The primary benefits of triploid plants are that they tend togrow larger and faster than In or 2n plants and that they aresterile. The ability to induce sterility in a plant will be a great
benefit to seaweed mariculture, since the plant would reproduce
vegetatively and could be cloned indefinitely. In addition to thevalue of the improved strains for cultivation, the basic
knowledge and other developments in this study should be applicable to other economic seaweeds; e.g., the use of polyploidy
to produce sterile clones of annual seaweeds, where maintaining
viable seed stock throughout the year is a major problem.
One final development of Chondrus cultivation is the recently
perfected isolation and axenic culture techniques by Chen andTaylor (1978) for medullary tissue explants from female
gametophytic plants of Irish moss; Chen and Taylor were able toculture the colorless medullary filaments of C. crispus, producing fronds with the normal morphology of the parent plant.
This is the first published report of successful preparation of
axenic tissue culture of a multicellular alga. Such cultures could

plastic. The seawater is fed at controlled rates with agitation
provided by blowing air through the tanks. In the future, warm
water from a nearby fossil fuel electric plant may be used in
the tanks to grow seaweeds at optimum temperatures yearround.
Other pilot cultivation projects with Irish moss were recently
underway in New York and Nova Scotia (Genu Products
Canada, Ltd.) A preliminary account of an experimental farming program at Montauk, N.Y., which involved a landbased
network of 15 reinforced-concrete tanks placed at descendingheights, was given by Moeller et al. (1973). This employed agreenhouse to heat the networks by solar energy and boost yearround production. Detached populations of Irish moss weregrown in the tanks to assay their yields of carrageenan, growth
rates, viability, and cleanliness. The cultivation facility of Genu
Products Canada, Ltd., in Nova Scotia used many of thebiological and physical parameters developed by the National
Research Council at Sandy Cove (Neish and Fox 1971; Neishand Shacklock 1971; Neish et al. 1977), but little specific information was available.
Several Canadian scientists recently showed that kappa carrageenan was restricted to the gametophytic plants of Irish
moss, while lambda carrageenan was present in the sporophytic
plants (Chen et al. 1973; McCandless et al. 1973; McCandless1974). Similar results were reported from natural populations bycomparing the differential gelling potentials and the presence orabsence of 3,6-anhydrogalactose, respectively in the gametophytes and sporophytes of C. crispus (Mathieson and 1\teter
1975; Craigie and Pringle 1978). As suggested by Shacklock andassociates (Shacklock, Robson, Forsyth, and Neish 1973;
Shacklock, Robson, and Simpson 1973) seed stock selection ofdifferential phases would provide a uniform and consistent supply of the different carrageenan fractions.
A Sea Grant project by D. Cheney and A. Mathieson at the
University of New Hampshire has attempted to apply genetic
studies to enhance cultivation of C. crispus through seed stock
selection and improvements in the plant. Specifically, the pro-32

".11 h uma

program. For example, Doty (1977) suggested that such introductions could aid in the stabilizing of supplies, the shortening of transport distances between farm and factory, and thepossible purification of wastewater discharges into the sea. Evenso, efforts to introduce new seaweeds in Hawaii currently have
involved 11 Federal, State, and local agencies, and similar efforts
in Guam required clearance from at least 7 Federal agencies. Dotyemphasized that new crop species should only be introduced afte r
careful consideration of possible damaging effects, but that theyshould not be strangled by bureaucratic complexities. he cited examples of the recent proposal for Macrocystis introduction to
Hawaii, as well as experiments with the introduction of the Tambalang strain of Eucheuma.
A considerable amount of information is available on
Eucheuma spp. found in Florida, including investigations of
their biology (Cheney and Dawes 1973, 1974a, b; Cheney 1975;
Cheney and Mathieson 1978b), ecology (Dawes, Mathieson, andCheney 1974), physiology (Mathieson and Dawes 1972, 1974;
Shipman 1974; Dawes 1976, 1979; Dawes et al. 1976; Moon andDawes 1976a, b; Dawes et al . 1978), and biochemistry (Dawes
1974b, 1977; Dawes, Lawrence, Cheney, and Mathieson 1974;
Shipman 1974; Cheney and Babbel 1975, 1978; LaClaire andDawes 1976). Much of this information is relevant to the plant's
mariculture and resource management. For example, the recent
electrophoretic studies of Eucheuma spp. (Cheney and Dawes
1973, 1974a, b; Cheney and Babbel 1975, 1978; Cheney 1976)
have obvious potential applications to both seed stock selection
and genetic improvement programs, as noted in the section on
Chondrus crispus.
A compilation of relevant data regarding the potential
mariculture of E. isijorme was summarized by Dawes (1974a),
who suggested that fragments could be dispersed and grown ineither natural embayments. which are common in Florida, or inculture tanks. In a natural habitat a number of factors should beconsidered to optimize the growth of E. isijorme (Dawes,
Mathieson, and Cheney 1974), including adequate water movement, water depth (0.5-3.0 m below ML W), fum substrata
without sand, and adequate illumination. Plants grown on lines(1974). Successful farming techniques evolved after a variety ofdifferent propagation methods, growth studies, and biochemicaland environmental investigations (Kraft 1970; Doty and Santos
1978). For example, constant-level planting techniques weredeveloped because of extensive grazing by fish and sea urchins(Doty 1973a, b; Parker 1974). Basically, about 10 kg of
Eucheuma are planted on a single net, with the nets being arranged in four modules with 800 nets each. The cost of installingand operating a 1-ha farm in the northern Sulu Sea area of thePhilippines is about US$3,126.00/yr, based on a 3-yr amortization. It is estimated that about 10 tons of dry weed per hectare
are produced, leaving 50-100 g of live weight of each thallus forseed. Large-scale Eucheuma farms seem to have the potential ofcompeting advantageously with other crops, including sugar andcoconuts (Doty 1973a, b).
There are four major problems associated with the increased
production (agronomy) of Eucheuma (Doty 1977): 1) The
danger of production monopolies in a developing country,
which may change its politics, policies, and regulations; 2) thetraditional advantage of merchants over individual farmers; 3)
the need to improve farming methods; and 4) governmental
bureaucracy and the management of introduced species. With
respect to the fust problem, Doty suggested that the best solution was to encourage farm development in several countries, including the U.S. Trust Territories of the Pacific Islands. In an
economic sense such competition would stabilize the price ofraw seaweed materials and improve the reliability of the supply.
The unreliability of the price of Eucheuma has discouraged
many farmers who cannot entrust their family's livelihood to a
wildly fluctuating market.
A major theme of the University of Hawaii's marine
agronomy program has been to enhance cooperation between
farmers, as well as to initiate better management, which would
increase and stabilize seaweed prices (Doty 1977). Different
methods of planting (lawn, hollow, and pen), as well as theselection of fast-growing plants such as the very successful Tambalang strain, were conducted (Doty and Alvarez 1975). The introduction of strains may also be advantageous to a mariculture

A concrete block used fo r outplanting studies of Florida Eucheuma by C. J . Dawes and associates in the Florida
Keys. Photograph by A. Mathieson.

34

detachment process itself. As noted previously, when young
sporelings were transplanted to the field, they became reproductively mature within 6-10 wk, which was similar to that ofaerated, detached macroscopic plants.
Temperature, rather than light, was the critical factor determining spore germination with G. tikvahiae, since it occurred in
darkness but not at low temperatures (Bird, McLachlan, andGrund 1977). Carpospores showed a lower percentage of germination than tetraspores, presumably because carpospores
were released simultaneously at various stages of maturity.
Tetraspores showed a continual release with an apparent highgermination percentage, which suggests they provide maximum
opportunity for spore germination. The process of fertilization
is not required for the production of tetraspores, which may beof considerable ecological importance.
A variety of field and laboratory studies on G. tikvahiae from
Prince Edward Island, Canada, have been summarized (Goldstein 1974a). Three actively growing forms were observed inestuaries and embayments on the Island: 1) Plants securely attached by a true holdfast to benthic bivalve molluscs; 2) detached,free-floating fronds; 3) large, bushy plants anchored in benthic
areas by byssus fibers produced by Mytilus (see also Goldsten
1974b). The latter form was very abundant in the late summer andwas found producing tetraspores and, to a lesser degree, carpospores. Both spore types were readily isolated in culture andsporeling development was studied. Goldstein (1977) also described the artificial recruitment and cultivation of G. tikvahiae in
Prince Edward Island. The tetraspores of the plants were seededon scallop shells in the laboratory, and after 1-2 wk of incubation
50,000 sporelings were outplanted at two sites in Cascumper Ba/
The outplants were monitored for 13 mo. The greatest decline insporelings, especially those incubated for only I wk, occurred during the rust month after outplanting, due to burial by sedimentation, grazing by herbivorous snails, epiphytism, and competition
for substrata with other animals and algae. Active growth ceased
when the water temperature fell below 15°C and the plants
entered a quiescent phase which was characterized by dark
pigmentation, brittle texture, and frond erosion. The plants remained quiescent from late September to mid-June, after whichtime active growth resumed. Most of the plants were sexually
reproductive in August.
The recruitment of G. tikvahiae sporelings on ceramic tiles(11 x 11 cm), which were submerged weekly during the summersof 1975 and 1976, was evaluated by Busby and Goldstein (1977).
~fter 1 wk in the field, as well as 2 wk under laboratory conditIOns, G. tikvahiae sporelings were counted on the tiles. High
sporeling density was observed in late July, concomitant with anabundance of in situ tetrasporic plants. A significantly lower
number of sporelings was noted during 1976 than 1975 coincident with lower light intensities and water temperatures.' A welldefined "edge effect" was observed on the tiles. Overall the tiles
appeared to have a substantial potential for augmenting recruitment of G. tikvahiae.
The growth of G. tikvahiae under greenhouse conditions(Neish and Fox 1971) has been determined (Edelstein et al.
1976). As would be expected from its natural distribution (Bird
Edelstein, and McLachlan 1977a, b), the plant's growth in tank~
was depe~dent upon warm-water temperatures (i.e., i 5 °_28 0C).
Equally lmportant was a proper combination of light andtemperature. The highest yield was an increase of 423Ofo /in o or
14.1 %/d at 20°C and 12-12 LD (light-dark) period.
Gametophytes showed optimal growth with 18 h of artificial il-

winter-germinated spores. Unfortunately the project was
discontinued in 1977 because of problems of maintaining seedstock throughout the year.

Graci/aria
Several Canadian scientists from the northeastern Atlantic
have recently initiated a variety of biological (Goldstein 1974a;
Bird, Edelstein, and McLachlan 1977a; Busby and Goldstein
1977; McLachlan and Edelstein 1977), taxonomic (Chapman etal. 1977; McLachlan 1979), ecological (Bird 1974-76, 1975;
Edelstein et al. 1974; Goldstein 1974a, 1977; Taylor 1975a; Bird,
Edelstein, and McLachlan 1977b; Busby and Goldstein 1977),
genetical (Meer 1977; Meer and Bird 1977; Meer and Todd
1977; McLachlan et al. 1977), chemical (Hong et al. 1968;
Duckworth et al. 1971; Young 1973, 1974), and cultural Edelstein et al. 1974; Goldstein 1974b, 1977; Bird 1975; Bird,
McLachlan, and Grund 1977; Edelstein 1977; McLachlan and
Edelstein 1977; Bird et al. 1979) studies of G. tikvahiae from the
Canadian Maritime provinces. ' Overall, their efforts are
directed to an increased utilization of Canadian agarophytes,
because of the growing commercial needs for agar (Mathieson
1975; Moss 1977). Preliminary summaries of many of these efforts have been presented at the Canadian "Agar andAgarophyte Workshops" (Anonymous 1973b, 1974b, 1975).
The ecology of attached populations of G. tikvahiae at Barrachois Harbour, Nova Scotia, was described by Bird (1974-76).
The growing season of the plant, which is characterized by watertemperatures of about 20°C, is < 3 m. Accordingly, the plant isrestricted to shallow, warm-water embayments (Edelstein et al.
1967; Bird, Edelstein, and McLachlan 1977b). When sporelingsof Gracilaria were transplanted to the field, they became
reproductively mature within 6-10 wk (Bird 1974-76).
Tetraspores were released in June and carpospores between Julyand September. The mode of reproduction of attached anddetached populations was different. Attached populations
reproduced via spores and exhibited an alternation of generations(Bird 1974-76). In contrast, free-floating populations, which wereusually dominated by tetrasporophytes, primarily perennated by regeneration of adventitious branches (Bird, Edelstein,
and McLachlan 1977a).
The in vitro life history of G. tikvahiae from the Canadian
Maritime provinces was recently described (Bird, McLachlan,
and Grund 1977). A typical Polysiphonia-type life history wasobserved in culture. The male to female ratio for plants
originating from tetraspores was 1: 1. In addition to a normal sex
ratio, the requirement of males for carposporophyte formationin culture provided presumptive evidence for the necessity offertilization to complete the life history. In contrast to earlier
studies of free-floating populations of Gracilaria where a scarcity of reproductive material was noted (Causey et al. 1946;
Stokke 1957; Kim 1970; Simonetti et al. 1970), detached plantsof G. tikvahiae showed an enhanced potential for reproduction
if aeration was applied. That is, the reproductive maturation ofthese detached and aerated plants occurred in 8-10 wk versus 8
mo for nonaerated cultures. The rapid induction of fertility indetached plants was attributed to favorable growth conditions(greater availability of nutrients and dissolved gases) and the'The species designated as Graci/aria tikvahiae from the northeastern coast of thenorth Atlantic has previously been designated as Graci/aria species or incorrectly
referred to as G. fo/ii/era (see McLachlan 1979).

36

plants weighing about 5 g each were fastened to vertical lines.
The plants were attached at seven depths, which were constantat all stages of the tide. Initially the plants showed a lag ingrowth for 1-4 d, presumably because of a change in light intensity after having been transplanted. The growth rate of the experimental plants was correlated with light intensity, with the
maximum growth occurring nearest the surface (0.3 ft), eventhough the plants were bleached. A comparison of the growthstudies in Florida and North Carolina showed that plants which
initially weighed 5 g increased 14-fold in 20 d at Alligator Harbor, Fla., versus 15-fold in Beaufort Harbor, N.C.
.
The growth rate of G. folii/era populations from North
Carolina, which were under approximately optimal conditions,
was only about one-half to two-thirds that of G. verrucosa
under the same conditions (Causey et al. 1946; Kim and Humm
1965). Both species appear to be very tolerant of high light intensities, as their maximum growth occurs near the surface of
the water; in addition the optimum growth for both is close to
the maximum temperatures of inshore waters of thesoutheastern' coast of the United States (28 °-32 0q. The
temperature tolerances of the two species seem to vary, as G.
folii/era is in good vegetative condition year-round throughout
its geographical range, while G. verrucosa is absent or dormant
during the winter north of Florida (Causey et al. 1946).
The physiological ecology of estuarine populations of G. verrucosa from Florida was recently studied (Dawes 1976; Dawes etal. 1978). The plant exhibited an increase in net photosynthesis
from 50 fc (1,150 /kW/cm2) to 2,000 fc (13,800 /kW /cm 2) : aswell as maximum net photosynthesis at 24 °_36 °C (30 °C optimum) and 10-50 '100 (30 '100 optimum). Accordingly, G. verrucosa shows a broad tolerance to light, temperature, and salinity, characteristic of its estuarine-mangrove habitat. The effects
of light quality on ammonium uptake with Floridian populations of G. verrucosa have also been described (Bird 1979). In
nitrogen-enriched cultures there was an enhanced ammonium
uptake under blue light, as compared to green or red light. In
contrast, nitrogen-starved cultures of G. verrucosa showed noeffect of light quality on ammonium uptake.
Additional investigations of Gracilaria from the Pacific coastof North America have recently been conducted. For example,
Whyte and Englar (1976a) evaluated the growth and chemicalcomposition of Gracilaria (= G. verrucosa?) in culture, based
upon British Columbia populations. The plants were propagated as free-floating entities in tanks which were flushed withfresh seawater and with effluent seawater originating from a fish
rearing tank. After a 113-d culture period, the plants from the
fresh seawater tank increased in mass by 21.3%. The plantsfrom the effluent seawater tank showed a 1.37% growth/dand an increase in mass of 55% after 113 d. Cultured specimens
had higher contents of dry matter and higher yields of agar thanin situ populations. The phycocolloid yields and properties ofother British Columbia populations of Gracilaria sp. have also
been evaluated by Whyte and Englar (l979a, b, 1980).
The growth and productivity of Graci/aria ( = G. verrucosa?)
from British Columbia, Canada, has also been studied(Saunders and Lindsay 1979). The reproductive strategy,
biomass production, and agar quality and quantity of intertidal
and subtidal populations were different. Regrowth in vegetatively propagating intertidal beds was more seriously affected thanin subtidal beds, which rely on spore recruitment. Culture
studies of free-floating populations were conducted, employing
six different phenotypes. A series of floating net enclosures were

Various approaches to culturing G. tikvahiae in tanks are being evaluated (Brochu 1977), including using plants with threesets of chromosomes or triploids. Triploid plants seem to belarger and grow faster than the normal (In or 2n) ones. Theiruse, along with other refmements in culture conditions, mightmake possible the commercial culture of G. tikvahiae in tanks orsalt ponds.
Several other studies dealing with G. tikvahiae from theAtlantic coast of the United States have recently been conducted. Fralick (1974) summarized the growth of G. tikvahiae(as G. folii/era in Taylor 1957) as well as Chondrus crispus andNeoagardhiella baileyi, in a nutrient-enriched, multipurpose
aquaculture system at the Woods Hole Oceanographic
Institute's Environmental System Laboratory. Specific details
regarding the Environmental System Laboratory are given in thesection dealing with Polyculture systems. Growth rates as highas 4O%/d were recorded, as well as the successful removal ofnutrients from the system. Further descriptions of the growth of
G. tikvahiae in the Environmental System Laboratory were
given by Fralick et al. (1975), DeBoer (1979), and LaPointe andRyther (1979). In addition, the net photosynthetic responses ofthe plant under a variety of different light, temperature, andsalinity conditions were described by Fralick and DeBoer (1977).
The optimal conditions for growth are 800-2,500 fc(footcandle), salinities of < 32 '100 and temperatures of
20°-30°C. The experimental results were interpreted in relation
to seasonal variations in growth.
The seasonal changes in chemical constituents and reproductive generations for G. tikvahiae populations (designated as G.
folii/era) from Great Bay, N.H., were enumerated by Penniman(1977). There were slight differences in agar yields between
vegetative, cystocarpic, and tetrasporic samples. Agar yields wereat a maximum from November to January, while protein andphosphate contents showed no consistent differences with season
nor reproductive status. The maximum occurrence of cystocarpic
and tetrasporic plants was in June and July, with cystocarpic
plants showing their maximum somewhat earlier thantetrasporic plants. The innermost estuarine sites exhibited earlier
reproductive maxima. Low percentages of cystocarpic plants
were consistently found at all three sites. Tetrasporic plants wereabsent in the innermost estuarine sites during the winter.
An additional study of biochemical properties of G. tikvahiae
was recently conducted in Rhode Island by Asare (1979a) inorder to optimize its harvesting and agar yields; he found thattissue nitrogen was highest during the winter. In addition, heemphasized that factors apart from nitrogen status were involvedin the increasing and decreasing of phycocolloid content in theseplants.
The agar variations of G. bursapastoris and G. coronopi/olia
from Hawaii have been reported by Hoyle (l978a, b). Crude
agar extracts from tetrasporic, male and female plants showedno significant differences in yield nor gel strengths (Hoyle
1978a), similar to that observed for G. tikvahiae (Penniman1977) The thallus nitrogen content of both Hawaiian species of
GraCI/aria was correlated with NO,-N in the seawater (Hoyle
1978b) In addition, the agar yields of both plants were inversely
related to the total nitrogen of the thalli. The reproductive
phenology and growth rates of G. bursapastoris and G. coronopifo/ta from Hawaii were also reported by Hoyle (l978c).
The ummer growth potential of G. folii/era populations at
Alligator Harbor on the Florida Gulf coast and at Beaufort,.c.. \\t:re determined by Kim and Humm (1965). Portions of38

de igncd and t ted in order to del ermineof depth. den II ,and phenol pe, \10 Ih Ihe
rowing Irain. I urther e penmenl I
lu'vera] . rra ccnoph)te , \IoL'TC conducled In r
ult ure condition In order tll rccn a lar number ot pI n
fa!t· I ll\lo 10 ' " tram .' (Buntin ct al. lit ).
The efle t of lighl inten ity and temperature on the rO\lo1hrate of cult urL'll G. n'rroco\o Irom \Va' hillgton VI ere rl'flOrtedby (an and \Vaaland (1977). 1 he 111 h t gro 11h rut
ere
recorded at 2!l.2"1oo • 28
and 1,400 fe. In general, the fO\Io1hrale\ in!;rea cd with increasing light inten lIle at aJltempcrature .
t ,aryin' light mten itie~, the gro\loth rate either de rcased or remained unchanged, C PL'(iallya temperature meretrom 2 0
to 2M • Only at 1.4(X) f wa\ there a ignific3nt Inlr
In
growth rate abm.e 23 c •In addition to the refercnce~ di~cu~scd abovc, the readerl>ho uld consult thc extensivc revie\lo of Gracr{arta gl\Cn by Hoyle(197 ). He summarile~ information regarding the ta onom).
orphology, ecology, phylogeny, and cconomic importance ofGractfarta. much of which IS relevant to the future manculture
and resource management of this plant. furthcr dcscriptions ofgrowth of Gractfaria in w~tc-rccycling polyculture s 'stem~ atWoods Hole, Mass., and in Florida (LaPointe ct aI. 1976; DeBoer
and Ryther 1977; Hanisak et aI. 1979), as well a~ In Rhode"land (Harlin et aI. 1979; A~are 1980), are givcn in the
Polyculture section.

Ilypl/ea
The red alga H~'Pnea is a species of considerable c\Conomlc\alue, as its cell \vall contains the poly\aceharide \.:apna \C'arrugeenan ( toUion 1973. 1979; Humrn and Kreule 19 5). At
present wild st cks arc insufficient to meet the requIrement of
the carrageenan produ(;ers; thus its mariculture ma) become
profitable \Iohen large· scale cultivation procedures are perfected .
e'eral recent studies ,ummarllC the sea anal 6rowth (Hummand !\:reulcr 1975; Mshlgeni 1977; hhlgem and Mliray 1979;
IIUIllI11 1911 I), reproduction ( 1shlgem 1976c). sporcllng dc\ elopl11t:nt (\tslllgcni 1976a. b. c). bilXhc:mistl) (\!shlgeni 1979;
Durako J.mJ DJ\\(; 19Sfu). ,l.J1d ph) ioloSY (DJ\\l: 1')"6: [).l\\~ et011 1976; Durako and Da\\e . 19 Ob) of HV[lflco. mu~h of \\hi.:h" rele\ ant to the future mariculturc of thi ~pecIC. An c tcnd~'(jrt:\ie\\ or the literature l1n HY[lfIlo 1\ al 0 a\"aIlablc (i\t hi em1974).

The
up\loclhn

the outdoor

The marine red alga Iridaea cordata has a very high carrageenan content (52-66070) and is of considerable economic
potential (Mumford 1977b; Waaland 1977). Several investigators have studied Iridaea evaluating the plant's growth(Fralick 1971; Waaland 1972, 1973a, b, 1974b, 1976,1977,1979,
In press; Hansen 1974, 1975, 1976, 1977a; Hansen and Doyle
1976; Adams and Austin 1977; Packard 1977), development(Norris and Kim 1972), reproduction (Norris and Kim 1972;
Waaland 1972, 1973a, b, 1974b, 1977, 1979; Hansen 1974, 1975,
1976; Hansen and Doyle 1976), cytology (Fralick and Cole1973), physiology-biochemistry (Waaland 1973a, b, 1974a,
1975; McCandless et al. 1975; Hansen 1977b), ecology (Fralick
1971; Waaland 1972, 1973a, b, 1974b; Austin et al. 1973; Austin
and Adams 1974; Hansen 1974,1975,1976, 1977a; Hruby 1974,
1975, 1976; Foster 1975b; Hansen and Doyle 1976; Adams andAustin 1977; Packard 1977), biogeography (Kim and Norris
1974; Kim 1976), and taxonomy (Abbott 1971,1972; Kim 1976).
Much of this information should be helpful in furthering themariculture and conservation of Iridaea.
Detailed accounts of the successful seeding of nets with carpospores and tetras pores of 1. cordata, as well as their subsequent yields and growth rates in various locations in Puget
Sound and the San Juan Islands, Wash., have been given (Kempand Mumford 1976; Mumford 1977b). Three types of nettings
were evaluated: A white type-66 nylon seine netting made of no.
60 cord, black plastic Vexar 6 (Dupont, Inc.), and orange
polypropylene. The plant's growth on the expanded nylon andpolypropylene netting was excellent, while the plastic Vexar was
unsuitable. The density of the plants on the best netting was approximately 10 times that in natural populations in Washington(Fralick 1971; Waaland 1976) and British Columbia (Austin andAdams 1974) and approximately 1.5 times that in central
California (Hansen 1976). The location and season of net
emplacement were found to be very important. For example, in1975 only about one-half of the nets placed were seeded withenough I. cordata to produce 100% cover the followi ng year.

There was no appre(.;iable increa\e in cover from additional
seedings during the second and third year~, due to coloni/ation
by ulvoids and kelp\. 1 hus, artificial \ecding of net~ In tanb
may be reqUired As the tetrasporic rlants of I. cordata contain
only lambda carrageenan and the gametophyul rlants only kappa carrageenan (Waaland 1974a, 1977; Mc andless et al. 1975),
it is possible to seed for one phase of the life history and get aparticular carrageenan. Two harvests arc ros ible rer year ""iththe fir t occurnng In . pring and the ~econd dunng Augu t oreptember when the maximum tanding crop occurs.
FertilizatIOn technique (e.g., ammonia) for rope-cultured
populatlOm of I. cordata are being perfected in order to determine their effects on growth, phenology, and carrageenan content (Mumford 1977b). 0 major diseases were recorded onrope outplants of I . cordata, although everal herbivores cau edextensive damage. Thu , Mumford (I977b) concluded that the
major obstacles to any aquaculture program, including I. cordata, may be ol:lal and legal, rather than biologic-d!,technological, or economical. Ho\\ever, if the u e of artifi ial
substrata for the culture of ea""eeds continue In Puget ound,
he suggested that the current te hniques could open ur ",ideareas of the 0 ean for production of hydrocarbons, energy,
food, chemicals, and pharmaceutical product. Further de riptlOns of I. cordata farming in the Pacific Northwest ha\e been
given (Jamison and Be wick 1972; Kemp and ~1umford 1976;\1umford I977a, 1979: Anon)mouS 197 '; lumford el al.
1979), including a hi torical a count of re earch pon ored by
the Washington tatc D\!partment of. atura] Re ource and the
Washington Sea Grant Program.
Growth experiments "'lth free-noaung populations of I. cordata, as well as Glgartma ewsperala, ha\e been condu ted insemiclosed culture system~, con Isting of large tanks supplied",,;th running ~ea\\ater (Wa'uand 19 6, 197 , 19 9, In pres ;Cooper 1981). He used evenl tank designs, y,hich \\ere baslcalJ}similar to tho e employed by:-'larine oUoid, Inc., for Chondrus
crispus and Eucheuma lSijorme (Deveau and astle 19 6.) The
major factor regulating the grO\\1h and optimal production of I.
cordata were plant size, rate of sea\\ater 110\\, ratio of blomas tosurface area, and light Intensity. The yield br I. ~ordala in uch aculture system was estimated as 5,700 kcal f m' per}T or 1,950 g
ash-free dry matter m' per yr, which was si\fold that of natural
populations (Waaland 1976). A companson of the annual production of 1. cordata with some major terrestial plants indicatedthat it compared favorably with the biomas produced from thehighest yielding corn, wheat, and rice farmmg technique(Waaland In press). A comparative summar} of the ummer
growth rates (as percent dry weight per day) of 13 Pacific orthwest marine algae showed that I. cordata and G. e'(asperata
were ranked first (9.5%) and fourth (8.31T'0), \\;th Porphyra perjorata (8.8%), I. cornucopiae (8.7%), and I. heterocarpa(8.1 %) showing comparable growth rates (Waaland 1977).
A major requirement for a sustained sea\\ eed aquaculture
operation, such as with I. cordata and G. exasperata, is theavailability of sufficient inocula each spring, prior to the major
growing season (Waaland 1977). In the case of G. exasperata
this is not a problem, as its large perennial blades are present
throughout the year, and they produce new blades from surface
and marginal proliferations. In contrast, I. cordata is a
pseudoperennial species, which has a perennial holdfast and'Reference to trade names does nOI imply endorsement by the National Marine
Fisheries Service, NOAA.

'Anonymous. 1977. Seaweed farming in the Pacific nonhwesl. Aquaculture
Newsletter #34, p. 1-3. Jerden Industries. Inc., Bellevue, Wash.

salinity between estuarine and coastal populations were noted,
which might be relevant to potential seed stock selection of fa tgrowing strains. Such methods can be employed to assay thephysiological state as well as strain tolerances to environmental
paraIJ1eters. Further descriptions of the physiological ecology of
H. muscijormis are given by Dawes (1976) and Durako andDawes (1980b) .
A model for the growth of detached H. muscijormis, based
upon the extensive outdoor cultivation program of the Marine
Colloids, Inc., at Summerland Key, Fla., from January 1975 toMarch 1976, was also summarized by Guist et al (1977). The
culture system consisted of large tanks with running seawater,
and was basically similar to that used by Marine Colloids, Inc.,
for C. crispus and Eucheuma isijorme (Deveau and Castle1976). The model related growth under a wide range of ambient
temperatures and light intensities to concurrent laboratory
measurements of the photosynthetic and respiratory activity of
H. muscijormis. The model allowed a more quantitative determination (assay) of the effects of water temperature and illumination on the growth rate of this species in culture.

Iridaea-Gigartina

40

blades that die off each year following reproduction. Even so, if
net cultures of I. cordata were established , they could produce
new blades each year from the perennial holdfasts (Mumford etal. 1979). According to Waalarid (1977), the blades of I. cordata
frequently produce spores 1-2 mo earlier in culture than in situ;
no such observations have been made with C. exasperata. One
particularly fast-growing strain (m-II) of C. exasperata wasmaintained by Waaland through three winters in outdoor
cultivation tanks. Two major factors made this possible: The
plant can be propagated vegetatively by means of cuttings oroutgrowths from the blades; and the plant can be sustained
through the winter, when the solar radiation is at a minimum,by adjusting the plant density per tank . Unfortunately, the sameprocedures cannot be followed with I. cordata, and in tankcultures the spring seed material must be initiated either by
juvenile sporelings (i.e., new plants), by the perennial holdfast
system, or by artificial substrata inoculated in the field. Thus,
Waaland (1977) suggested that C. exasperata was a better candidate for tank mariculture than I. cordata, especially if strains
with faster growth rates and yields were developed. In this vein,
Waaland (1977, 1979) has conducted some strain selection
studies by growing a variety of plants under uniform conditionsfor 4-6 wk periods. He found that fast-growing strains can beisolated, propagated by vegetative means, maintained insemi closed culture for at least 2 yr, and grown in large quantities.
For example, the m-II strain increased its fresh weight at a rateof 4.25 OJo/d versus 3.II% / d for wild material.
A variety of productivity studies of I. cordata have been
described (Hansen 1977b). The time periods required for a
doubling in dry weight, as well as carbon and nitrogen contents,
were used to evaluate the gap between laboratory-determined
and in situ estimates of productivity as well as to identify
limiting parameters. During the winter, spring, and summer, insitu weight and thallus carbon doubling times were similar, indicating that laboratory-determined photosynthetic rates, forthese seasons, were indicative of field produdion rates. During
the fall, when natural populations senesced , laboratorydetermined photosynthetic rates gave inaccurate estimates offield production. Doubling time comparisons of in situ populations suggested that production was light-limited during winterand spring but limited by other factors during summer andautumn. Laboratory-determined doubling times for thallus
nitrogen were longer than in situ times during all seasons except
winter. This suggested that in situ thalli utilized additional
nitrogen sources not considered in this study or that the thalli
carried out luxury consumption and stored nitrogen for utilization when seawater sources were depleted.
The photosynthetic rates for a fast-growing strain (m-II) of
G. exasperata were evaluated by Merrill and Waaland (1978,1979). The photosynthesis and respiration rates of the plants (i.e.,
blades) were measured and compared with values similarly obtained from wild material of the same species. Measurements
were made with an oxygen electrode over a wide range of lightintensities (5-750 !!E/m' per s) and temperatures (6°-22°C), in
order to predict the effects of manipulation of light intensities
and temperatures in aquaculture operations.
Genetic improvement studies of seaweeds may be critical in
expanding the production of certain limited and / or underutilized
economic red algae, such as the carrageenophyte Cigartina(J. West', In press). Thus, West and his associates have initiateda genetic program for the improvement of carrageenan production in Cigartina subgenus Mastocarpus, a kappa carrageenan
producing species of high quality. Most of their efforts havecentered on C. papillata and C. agardhii, which may exhibit twolife histories, a heteromorphic alternation of generations between a gametophytic Cigartina phase and a crustosetetra sporophyte (Petrocelis) or an apomictic recycling of Cigartina (West 1972, In press; West and Polanshek 1972, 1975, 1977;
Polanshek 1974, 1975; Polanshek and West 1975, 1976, 1977;
West et al. 1978). A major premise in West's efforts is that there
is a genetic control of carrageenan synthesis in several Gigartinalean red algae (Chen et al. 1973; Pickmere et al . 1973, 1975;
McCandless et al . 1975; Waaland 1975), with the primary typesof carrageenans being linked to the nuclear and morphologicalphases of the life history, i.e., gametophytes primarily produce
kappa carrageenan and tetrasporophytes produce mainly lambda carrageenan. Accordingly, a knowledge of the plant's life
history, whether it be apomictic or a heteromorphic alternationof generations, may enable a selection of specific carrageenans
via different reproductive generations or strains of Cigartina. At
present West is attempting to produce high-yielding strains of C.
papillata and C. agardhii. In addition, promising strains havebeen grown and crossed in the laboratory and their progenytransplanted into the field. Analyses are being conducted inorder to determine the types and quantity of carrageenans inboth the parents and progeny.
The influence of substrata and spore concentration on spore
survival and germination in C. agardhii and Petrocelis jranciscana has been evaluated (West and Crump 1974). The carpospores of C. agardhii and the tetraspores of P. jranciscana
were germinated on three different substrata: Microscope slides,
polystyrene petri dishes, and Plexiglas (acrylic) plates. Spores
liberated 24 h or more showed greater developmental abnormalities than freshly liberated ones (0-6 h old). A greaternumber of spores developed as discs on smooth polystyrene andglass than on Plexiglas. In addition, growth was more rapid on
polystyrene and glass than on Plexiglas. At the lowest concentration tested (1,080 spores/cm') P. jranciscana spores did notsurvive; however, at higher concentrations many spores germinated and developed as normal discs. With C. agardhii onpolystyrene, carpospore concentration had little apparent influence on the percentage germination. However, it did have adramatic effect on germling development. More carpospores of
C. argardhii developed as discs in low concentrations (63%)
than in high concentrations (9%). Only spherical germlings
developed on Plexiglas and about 10% were discoid on glass. Of
the substrata tested, Plexiglas was least suitable for outplanting,whereas polystyrene appeared best because it permitted rapid
spore germination and attachment of discs.
Additional information on Cigartina carpospore discharge
periodicity was summarized by West and Crump (1975). Eachpapilla discharged carpospores at 14(1O-16)-d intervals with
discharge occurring on 2-3 successive days during each cycle. In
addition, each papilla discharged 5,000-100,000 spores/d. Thus,
the total spore production per papilla averaged 3,600,000.
A variety of studies have been conducted on C. stellata, aplant which is often harvested in mixed lots with C. crispus as asource of carrageenan (Burns and Mathieson 1972a). For example, the seasonal growth, reproduction, and ecology of naturaland harvested populations have been evaluated (Burns andMathieson 1972b). In addition, culture and morphological•John West, Botany Department, University of California, Berkeley, pers. commun. January 1975 .

41

were recently interpreted (Santelices 1978b). The plants'
biological cycles were signiflcantly correlated with seasonalchanges in light intensity and water movement but not withseasonal changes of water temperature and salinity. Laboratory
experiments tested the effects of five single factors and nine
types of interactions on plant growth. Water movement andlight intensity were the primary factors regulating growth. Water
enrichment compensated for water movement, as all three
species attained maximum growth at comparatively low water
movement intensities when grown in fertilizer-enriched media.
A preliminary account of an artiflcial flotation technique forthe cultivation of Gelidium sp. was summarized by Gibor(1976). He described a Simple and inexpensive system forfloating Gelidium plants near the surface of water with perlite
particles, i.e., aerated volcaniC rocks . Small fronds of Gelidium
were glued to the particles, while other plants such as Bryopsis
attached themselves. Gibor suggested that the potentials formass cultivation of Gelid/um using these methods were excellent.
One of the few papers dealing with ma<;.s culture of gelidioid
algae (i.e., P. caerulescens, P. capillacea, and Gelidiella acerosa)
is given by Santelices (I 976b). Working with Hawaiian material,
he evaluated the growth of unattached specimens as well as
minimum water volume reqUirement per unit of algal mas .
Gelldiella acerosa was unable to grow detached. However, P.
caerulescens and P cap/llacea grew continuously for 50 d,reaching maximum daily growth rates of 2.3 and 1.6070 reS'pectively. Both of the latter species adopted a globose or subglobo e
habit, showing modifications similar to other free-floating
species of algae (Austill 1960). The optimum \olume per unit ofalgal mass was 1 liter of enriched seawater per 3.3 g for P.
caerulescens and 1.7 g for P. capillacea.

studies were made to determine the plant's life history (Chen etal. 1974; Edelstein et al. 1974; TVeter and Mathieson 1976a), andgrowth and ecology (Burns and Mathieson 1972a). The
physiological ecology (Mathieson and Burns 1971) and carrageenan ecology of G. stellata (Mathieson and TVeter 1974,
197~ have also been investigated. Abbott (1980) has recorded
similar evaluations of colloid variations in several red algae incentral California, including the kappa-carrageenan producing
species Gigartina leptorhyncos.

Gelidium-Pterocladia
Extensive reviews of the gelidioid algae, including their taxonomy, morphology, ecology, phylogeny, and economic importance, have been summarized (Santelices 1974, 1976a, 1977,
1978a, b). Most of the agronomy-related studies of this group
have been oriented towards improving harvests and protecting
the species as a natural resource (Santelices 1974). Barilotti andSilverthorne (1972) gave an excellent evaluation of resource
management of subtidal populations of G. robustum insouthern California, based upon a variety of ecological information including biomass and agar variations, growth, reproduction, regeneration capacity, and seasonal and habitat differences(Cooper and Johnstone 1944; Johnstone and Feeney 1944;
Barilotti 1974). Silverthorne (1973, 1977) also developed aharvesting model for G. robustum based upon the above information. According to this model, a natural population has astanding crop of about 500 glm' (fresh weight). After harvesting
and regrowth for 14 quarters, the standing crop would be 900
glm' and it could be reharvested with minimal effects. The
harvesting model permits a prediction of yields under various
management schemes.
The ecology, distribution, and abundance of G. robustum insouthern California was described by Barilotti (1974). He suggested that light, water motion, and their interactions influenced
vegetative growth rates. Transplant studies indicated that water
motion, through its effect of limiting grazing pressure, wasresponsible for determining frond density. Barilotti also suggested that the disappearance of G. robustum from large areasof the California coastline was due to pollution.
The growth, reproduction, and harvesting of the commerically important agarophyte G. robustum (as G. cartilagineum var.
robustum) in Baja California has also been described (Guzmandel Pro'o and de la Campa-de Guzman 1969). The growth of
harvested and nonharvested (control) plants was assessed, inorder to evaluate their regrowth potentials. Tetrasporic plants
were predominant throughout the year, while carposporic plants
were observed in smaller numbers and for more limited periods.
The spore dispersal of the plants was found to be predominantly
southeast from the parent populations. Spore shedding and germination of G. robustum was also studied by Guzman-delPrdo et al. (1972). The plants showed a maximum reproductive
activity during August and September, and minimal activity in
January and February. Both tetrasporic and carposporic plants
showed diurnal discharge periodicities; in each case, carposporicplants released the greatest number of spores. Guzma'n-del
Pro'o and de la Campa-de Guzman (1979) further summarized
a variety of synoptic, ecological, and biological information for
G. robustum populations in Baja California.
The multiple interaction of several factors in deflning the
distribution and abundance of three Hawaiian Gelidiales(Gelidiella acerosa, Pterocladia caerulescens, and P. capillacea)

Little is known about the nutrient physiology of seaweeds, including that of the gelidioid algae. Bird (1976a, b) evaluated thephysiological responses of the agarophyte Gelidium nudifrons
to various combinations of ammonia and nitrate. He observed asimultaneous assimilation of both nitrogen sources in culture experiments, employing four different combinations of the twonutrients. His study suggested that there were pools of bothnutrients which were simultaneously available for algal assimilation.

Palmaria palmata (Dulse)
An appraisal of the dulse industry in Canada, including ahistorical analysis of the industry, an evaluation of the resource,
and its marketing and developmental opportunities, was recently given by Ffrench (1974). Dulse is usually harvested by hand(rake), air dried and packaged as a source of food (Morgan et al.
1980; Sirota and Uthe 1979). One company, Atlantic Mariculture Ltd. of Grand Manan Island, New Brunswick, Canada,
has utilized impoundment and cultivation techniques (Anonymous 1974a; Neish 1976a), which consisted of ferrocementtanks and air agitation. The company held detached plants intanks until they were processed; any growth that occurred offset
the operating cost of impoundment. Neish (1976a) stated that
dulse harvesting is only carried out during very low tides ofspring and fall, so the flow of raw material thro~gh a processing
facility is uneven. A capacity for holding live material greatly
enhances processing efficiency. Dulse is cleansed of dirt,
mussels, epiphytes, and other contaminants in the storage tanks.

42ie as of overg ry wt7m per a were 05taine witli stocKing
densities of approximately 10 kg fresh wt/ m'. He suggests that
P. pa/mata, which has an annual frond and a perennial basal
system, must be propagated differently than perennial species
like Chondrus crispus. Such techniques may require thedevelopment of new methods involving grow-out from hatcheryseed. Some commercially significant marine plants have beensuccessfully brought through their life-cycle in the laboratory,
and tht;! commerical production of hatchery seed is wellestablished in the Porphyra industry in Japan (MacFarlane
1968; Okazaki 1971; Korringa 1976).
The effects of total carbon supply, irradiance, pH,temperature, and salinity on the short-term photosynthetic ratesof P. pa/mata were enumerated by Robbins (1979), based upon
oxygen and pH measurements. Photosynthetic, but not
respiratory; rates showed some seasonal adaptation. Maximal
photosynthesis occurred at 10 °-15 °C, 32 '7'00' and pH values of
6.5-7.5. Light saturation was at approximately 212 jili/m' per s
and P max was temperature dependent. The compensation
point in late winter was 6.6 jili/ m' per s. Raising the total carbon dioxide concentration to 6.0-7.0 mM/ 1 resulted inphotosynthetic stimulation up to 2.7 times normal levels. A
photosynthetic quotient of about 1.28 was determined for
Pa/maria pa/mata by measuring carbon dioxide flux with infrared gas analysis.
The life history and cytology of P. pa/mata from the Atlantic
coast of Canada have recently been studied (Meer and Chen
1979). Tetraspores from diploid tetrasporophytes, the most obvious plants in situ, gave rise to two types of haploid sporelings,
one of which grew more robustly and was less branched than theother. AI: 1 ratio of the two types was obtained from random
spores. The more robust plants were males, while the smaller
plants, which were believed to be females, grew more slowly andeventually formed sporangia. The sporangia on the female
plants aborted without releasing viable spores, presumably
because of nuclear abnormalities associated with haploidmeiosis. On one occasion fertile (i.e., diploid) tetrasporic frondsof P. pa/mata were obtained from gametophytes in culture, andthese gave rise to a second generation of segregating sporelings.
As noted by Meer (1976) in an earlier cytological study of P.
pa/mata, a complete elucidation of the plant's life history will
only be achieved when it is determined how the diploid state isattained. Since diploid tetrasporic plants appear to give rise only
to haploid plants, a compensating mechanism such as fertilization or somatic chromosome doubling must exist to generate
diploid plants. Hopefully, this mechanism can be found by
future studies. An ultrastructural study by Pueschel (1979) has
also demonstrated that meiosis occurs during tetrasporogenesis
in P. pa/mata.

creased interest in oriental cookery by Americans as well as thepresence of large numbers of Japanese in the United States andCanada have also stimulated the markets for nori.

Porphyra is a potentially important food resource in the
United States, as approximately US$700,OOO worth of nori is imported through San Francisco alone (NeushuI 9 ). In addition, therate of consumption of nori is increasing in the United States.
As a result, Neushul and his associates (Woessner 1974, 1979;
Woessner et al. 1977) at the University of California, Santa Barbara, have initiated several biological and economic studies on
Porphyra nereocystis, in order to provide necessary information
for intelligent management and cultivation of this underutilized,
but potentially important, species. Neushul (footnote 9)
estimated that the natural crop within a 55-mi distance of the
California coastline could be worth in excess of US$500,OOO.
Extensive nori cultivation may be possible in Puget Soundbecause the area has moderate winter (6 °_8 0q and summer
temperatures (seldom > 15 0q, a lack of sea ice, high nutrients
due to runoff from nearby mountains, adequate currents, andsufficient light for the growth of many Porphyra species(Hunter and Nyegaard 1974). Because the amount of stable
substrata is limited in shallow water, artificial substrata would
have to be employed in order to use the great body of water in
Puget Sound for seaweed production. Commercial culture ofnori would also require a mechanical harvesting system andlocal processing of nori sheets.
One of the few accounts of mass cultivation of Porphyra in
North America (i.e., in Washington) is given by Waaland(1977). He compared the growth of free-floating populations ofP. perjorata in a semiclosed culture system with 12 other PacificNorthwest red algae, including the carrageenophytes Iridaea
cordata and Gigartina exasperata. Porphyra perjorata showed asummer growth rate of 8.8070 dry weight increase per day, with
I. cordata being the only species tested with a higher growth rate
(9.5%).
Sea Grant studies by M. B. Allen and C. P. McRoy at the
University of Alaska examined the feasibility of Porphyra
cultivation in Alaska (W. Shaw IO ). The studies included anevaluation of favorable sites for cultivation, a determination of
the time course of the alga's life history under Alaskan conditions, laboratory cultivation of germlings suitable for inoculation in favorable habitats, and the establishment of an experimental "farm."
Another Sea .Grant project by D. Kapraun of the Universityof North Carolina (Wilmington) was initiated in 1977, in orderto determine the feasibility of establishing a nori harvesting industry in North Carolina. He assessed the distribution and standing crop of Porphyra in North Carolina tidal marshes, determined harvesting and processing techniques, and initiated apilot production plant and marketing procedures for the local
product.

Porphyra (Nori)
Porphyra, or nori as it is commonly called, is extensively
cultivated in the Orient (Korringa 1976) and is one of the mostimportant economic seaweeds in the world (Mathieson 1975).
Imada et al. (1972) stated that the annual sales of nori in 1970
were about US$I40 million, which was higher than any othermarine product in Japan. As a result of increased pollution ofJapanese coastal waters (culture areas), as well as increased demand for the product, the Japanese are seeking additionalsources of foreign Porphyra. Accordingly there is considerable
potential for the development of culture systems of nori inNorth America (Hunter and Nyegaard 1974; Hunter 1975). In-

The nutritional composition (McLachlan et al. 1972), life
history (Bird et al. 1972; Bird 1973), and ecology (Bird 1973) of
P. linearis have recently been studied in Canada. McLachlan etal. (1972) determined that P. linearis was superior to many other'Michael Neushul, Biology Department, University of California, Santa Barbara, pers. commun. January 1975.
"William Shaw, NOAA Office of Sea Grant, Washington, D.C. , pers. commun .
March 1971.

43

Species of nori used in the Orient, "'ith its desirable flavor

resulting from a high concentration of isofloridoside and several
free amino acids. The life history of P. linearis was completed inulture through fi\e generations (Bird et al . 1972). Conho porangia formed at 5 _20 °C, but release of conchospores
began only at 13 °C and continued at a slower rate at 5°C. The
conchocehs phase was long lived despite continuous sporulation
and attrition. Variations m ability to shed conchospores indicated the existence of different strains of conchocelis. Additional details regarding the life history and ecology of in situ
populations of P. Imeans were summarized by Bird (1973).
A detailed account of sexual reproduction in P. gardneri wasrecently summarized by Hawkes (1978). He observed fertilization and diploid chromosome numbers in the carpospores andvegetative cells of the conchocelis phase, which was the first unequi\ocal e\ ence for sexual reproduction in Porphyra.
Ha\\ kes (1 977a, b) further summarized a variety of field,
culture, and cytological studies of P. gardneri.
The gro"'1h of detached P. umbilicalis in a flowing seawaterS} tern at the Manne Plant's Experimental Station near~liminegash, Prince Edward Island, Canada, was reported by
McDougall (1974). In one experiment, detached specimens gained 107010 weight (wet) during 42 d, showing that P. umbilicalis
could be successfully cultured as unattached plants. Several
small rocks bearing attached reproductive plants were also
cultured in shallo'" tanks (18 m deep) with running seawater.
The tanks ere drained each day, leaving the plants exposed toair for about 4 h, approximating the conditions prevailing in
nature. Porphyra umbilicalis grew well under these conditions,almost doubling in length in 42 d. Furthermore, small plants
de\eloped on the rocks, suggesting the potential use of various
ubstrata for the growth and transfer of sporelings as is done"'1th the Japanese nori industry (Mathieson 1975 ; Korringa1976).
A vanety of other studies dealing with the life histories(Hollenberg 1958; Kri hnamurthy 1969a, b; Bourne et al . 1970;
hen et al. 1970: Conway and Cole 1973; Mumford 1973a, b,
1975; Karlfaun and Luster 1980), taxonomy (Krishnamurthy
1% a, 1972, ~lumford 1973a, b, 1975; Conway 1975 ; Conwayet
a1. 1975), ecology (Edelstem and McLachlan 1966; Mumford
I 3a), and ultural perspective-significance (Williams 1979) ofI'orph~ro ha\ e been conducted inorth America. As noted
earher, u h ~tudle may ultimatel) aid in the utilization anddc\elopmem of uch marine plant re ouree .
0I aminaria

hapmanme ured pcn<Xilcwdll)
hoi
that
n the bl d . A thepunctures approached the tips of the blades, new ones weremade at the base. The blades behaved like moving belts oftissue, eroding at the tips while growing at the bases, so that a
year's growth amounted to 1-5 times the initial length. Surprisingly, growth was most rapid from January to April, i.e., during
periods of short day length, low light intensity, and low watertemperatures. Conversely, it was slowest during July to October, when contrasting environmental conditions occurred.
The authors suggest that during the winter the plants are using
some of the dry matter stored in the summer as an energy andmaterial source for part of their growth. Since winter is time ofhigh concentration of dissolved nitrogen and phosphorous in
the sea, it seems possible that the plants are making some of
their new tissue by using a stored carbon source and taking up
nutrients from the seawater. This could obviously place them inan advantageous position for making maximum use of sunlightfor photosynthesis in early spring.
The summer biomass of kelps in St. Margaret's Bay has a turnover of 4-10 times in the course of 1 yr (Marm I 972b). Marm
estimated thilt the annual production of kelps in St. Margaret's
Bay amounts to 603 g Cl m' averaged over the whole bay, 1,750
g C/m 2 averaged over the seaweed zone, or 648,000 g Cl m ofshoreline. The approximate caloric equivalents were 6,030
kcal/m 2 over the whole bay or 648 x 103 kcall m of shoreline.
Overall, the growth strategy of kelps seems to center aroundtheir ability to carry out intensive photosynthesis in the summer,
as well as to store and mobilize carbon reserves during periodsof low light intensity (Mann 1973; Mann and Chapman 1975).
Such a growth strategy would greatly extend their growing
season and allow the plants to use the nutrients for growth during a time (winter) when the phytoplankton show slow growth
due to the lack of an energy source.
The seasonal fluctuation in dissolved inorganic nutrients, internal reserves of nitrogen, and growth rates of L. longicruris in
St. Margaret's Bay were summarized by Chapman and Craigie(1977). The onset of winter growth, as noted by Mann (1972b),
correlated with improved nitrate conditions in the sea. During
the winter, tissue reserves of nitrate reached 150 j.tM /g fres hweight in March, which was'a concentration factor of approximately 28,000 over ambient (water) levels. Depletion of thisnutrient pool followed the disappearance of the external nitratewith a lag period of up to 2 mo. Rapid kelp growth wasmeasured at this time. Reserves of organic nitrogen also reachedmaximum values in March and declined slowly throughout thesummer into autumn . Accordingly Chapman and Craigie (1977)
suggested that the combined inorganic and organic nitrogen
reserves sustained rapid growth rates into July and reduced rates
through the late summer. Fertilization of an experimental kelp
bed with NaN0 3 increased the internal plant reserves of nitrate
and produced a much improved summer growth rate. The
enriched plants developed very small reserves of carbohyrate
dunng this period .
The role of carbohydrate reserves as a carbon source during
the seasonal growth of L. longicruris has been determined(Chapman and Craigie 1978) . Seasonal variations in the levels oflaminarin and mannitol showed maximum value during latesummer. Minimum values occurred in February at the time
when growth rate were increasing rapidly. Erosion of the bladesin ",inter carried away much of the tissue containing carbohydrate reserve accumulated during the previou summer.
penmental reducllon of carbohydrate reserves by fertilization",ith odium nitrate during the summer did not affect sub equent "'Inter growth rate~. Truncation of the plants inaNovember significantly reduced growth rate. The lower rate of
growth wa~ attributed to a reduction in photosynthetic arearather than a loss of storage product~. An additional de criptionof the annual carbon budget for L. lonRicruri5 in Sl. I 1argaret'~
Bay was given by Hatcher et al. (1977).
The seasonal patterns of nitrate uptake in L. longicruris were
determined by Harlin and Craigie (1976), based upon populations from St. Margaret's Bay. Their experiment suggested that
L. long/cruris was capable of using nitrate under winter conditions, and that it did not depend upon stored compounds for anitrogen source during periods of rapid growth, i.e., late winterand spring. Discs were punched from the thin edges of L.
longicruris plants and they were subjected to external nitrate in aseries of concentrations under different light and temperature
regimes. The optimal rate of uptake, with summer material(June and July) was 7-9 JLM /g dry wtlh at 15 °C and approximately 700 fc. At 5°C, uptake was 60%, and at O°C it was 40070
of that at 15°C. The uptake in the dark was 60-80070 of that in
the light. Algae pretreated for 12 h with antibiotics removed
nitrate from the medium at a rate of 84070 of the untreatedmaterial. Nitrate uptake was inhibited in proportion to the concentration of nitrate in the medium. Further studies of nitrate
uptake by L. longicruris have been reported by the same investigators (Harlin and Craigie 1978).
The growth and production of L. longicruris are interrelatedwith nutrients, water motion, and other parameters (Gerard andMann 1979). That is, measurements of internal inorganic
nitrogen (i .e., nitrite and nitrate) reserves and dry organic weightindicated that low concentrations of dissolved nutrients insummer and low levels of illumination in fall and winter weremore limiting to growth at exposed than sheltered sites.
The morphological differentiation of nondigitate Laminaria
populations from the North Atlantic was evaluated by Chapman (1973a, b), who found that the occurrence of long, hollow
stipes (the L. longicruris types) decreased with increasing
exposure. The differentiation of population~ in relation to
exposure was elinal with overlap in the variation patterns The
major taxonomic finding of the analyses was the lack ofnonoverlapping criteria by which L. agardhii and L. fong/crurlS
may be delimited. Subsequently, Chapman (l973b, 1974a) has
conducted heritability te ts between the two morphologicaltypes within the Simplices section of Laminaria. The productionof fertile hybrids e tablished interfertility between populations
from the extremes of the morphological range. Reciprocal
Iran plantation experiments between exposed and sheltered sites
produced no significant change in tipe morphology. Quantitative genetic analysi of the degree of re emblance between
relati es indicated a high genetic component in the Interpopulation differentiation. Estimate of within-population heritabiht)
at the exposed Ite were low, perhaps because of intensi\ e local
stabilizing election which reduce genetic \'ariabilit).
The Inheritance of mucilage canal in Laminar/a, section
implice , from eastern Canada has alo been de,cribed b)
Chapman (l974b, 1975). A matri, of heritability test. hO\\cd
complete mterfertihty between all mucilage anal type, femle
F. hybrids beIng produced m all a. e~. Quanlltati\ e geneticanal), is re\ ealed a large emlronmenta.l component in thephenotypic \ anance of degree of mucilage canal de\t~lopm nt
Onl~ plant. from 0\ a eOlIa bred true \\ irh rl.: pct to muul·anals. ThIS characteri,tlc \\ a. cconidered un uitabk f r t -in expenment In the n ndl II teboth Id~ 0 thl.:

punctures approached the tips of the blades, new ones weremade at the base. The blades behaved like moving belts oftissue, eroding at the tips while growing at the bases, so that a
year's growth amounted to 1-5 times the initial length . Surprisingly, growth was most rapid from January to April, i.e., during
periods of short day length, low light intensity, and low watertemperatures. Conversely, it was slowest during July to October, when contrasting environmental conditions occurred.
The authors suggest that during the winter the plants are using
some of the dry matter stored in the summer as an en:rgy andmaterial source for part of their growth. Since winter is a time ofhigh concentration of dissolved nitrogen and phosphorous in
the sea, it seems possible that the plants are making some ofthe;r new tissue by using a stored carbon source and taking up
nutrients from the seawater. This could obviously place them inan advantageous position for making maximum use of sunlightfor photosynthesis in early spring.
The summer biomass of kelps in St. Margaret's Bay has a turnover of 4-1q times in the course of I yr (Mann 1972b). Mannestimated that the annual production of kelps in St. Margaret's
Bay amounts to 603 g Cl m' averaged over the whole bay, 1,750
g Cl m ' averaged over the seaweed zone, or 648,000 g Clm of
shoreline. The approximate caloric equivalents were 6,030
kcall m' over the whole bay or 648 X 10' kcallm of shoreline.
Overall, the growth strategy of kelps seems to center aroundtheir ability to carry out intensive photosynthesis in the summer,
as well as to store and mobilize carbon reserves during periodsof low light intensity (Mann 1973; Mann and Chapman 1975).
Such a growth strategy would greatly extend their growing
season and allow the plants to use the nutrients for growth during a time (winter) when the phytoplankton show slow growth
due to the lack of an energy source.
The seasonal fluctuation in dissolved inorganic nutrients, internal reserves of nitrogen, and growth rates of L. longicruris in
St. Margaret's Bay were summarized by Chapman and Craigie(1977). The onset of winter growth, as noted by Mann (1972b),
correlated with improved nitrate conditions in the sea . During
the winter, tissue reserves of nitrate reached 150 j.tM/g fresh
weight in March, which was a concentration factor of approximately 28,000 over ambient (water) levels. Depletion of thisnutrient pool followed the disappearance of the external nitratewith a lag period of up to 2 mo. Rapid kelp growth wasmeasured at this time. Reserves of organic nitrogen also reachedmaximum values in March and declined slowly throughout thesummer into autumn. Accordingly Chapman and Craigie (1977)
suggested that the combined inorganic and organic nitrogen
reserves sustained rapid growth rates into July and reduced rates
through the late summer. Fertilization of an experimental kelp
bed with NaNO, increased the internal plant reserves of nitrate
and produced a much improved summer growth rate. The
enriched plants developed very small reserves of carbohyrate
during this period.
The role of carbohydrate reserves as a carbon source during
the seasonal growth of L. longicruris has been determined(Chapman and Craigie 1978). Seasonal variations in the levels oflaminarin and mannitol showed maximum values during latesummer . Minimum values occurred in February at the time
when growth rates were increasing rapidly. Erosion of the blades
in winter carried away much of the tissue containing carbohydrate reserves accumulated during the previous surrUner.
Experimental reduction of carbohydrate reserves by fertilization
with sodium nitrate during the summer did not affect subsequent winter growth rates. Truncation of the plants inspecies of nori used in the Orient, with its desirable flavor
resulting from a high concentration of isofloridoside and several
free amino acids. The life history of P. linearis was completed inculture through five generations (Bird et al. 1972). Conchosporangia formed at 5 0 -20°C, but release of conchospores
began only at 13 °C and continued at a slower rate at 5°C. The
conchocelis phase was long lived despite continuous sporulation
and attrition. Variations in ability to shed conchospores indicated the existence of different strains of conchocelis. Additional details regarding the life history and ecology of in situ
populations of P. linearis were summarized by Bird (1973) .
A detailed account of sexual reproduction in P. gardneri wasrecently summarized by Hawkes (1978). He observed fertilization and diploid chromosome numbers in the carpospores andvegetative cells of the conchocelis phase, which was the first unequivocal evidence for sexual reproduction in Porphyra.
Hawkes (1977a, b) further summarized a variety of field ,culture, and cytological studies of P. gardneri.
The growth of detached P. umbilicalis in a flowing seawater
system at the Marine Plant's Experimental Station near
Miminegash, Prince Edward Island, Canada, was reported by
McDougall (1974). In one experiment, detached specimens gained 1070/0 weight (wet) during 42 d, showing that P. umbilicalis
could be successfully cultured as unattached plants. Several
small rocks bearing attached reproductive plants were also
cultured in shallow tanks (18 in deep) with running seawater.
The tanks were drained each day, leaving the plants exposed toair fo r about 4 h, approximating the conditions prevailing in
nature. Porphyra umbilicalis grew well under these conditions,almost doubling in length in 42 d. Furthermore, small plants
developed on the rocks, suggesting the potential use of various
substrata for the growth and transfer of sporelings as is donewith the Japanese nori industry (Mathieson 1975; Korringa1976).
A variety of other studies dealing with the life histories(Hollenberg 1958; Krishnamurthy 1969a, b; Bourne et al. 1970;
Chen et al. 1970; Conway and Cole 1973; Mumford 1973a, b,
1975 ; Kapraun and Luster 1980), taxonomy (Krishnamurthy
1969a, 1972; Mumford 1973a, b, 1975; Conway 1975; Conway etal. 1975), ecology (Edelstein and McLachlan 1966; Mumford
1973a), and cultural perspective-significance (Williams 1979) ofPorphyra have been conducted in North America. As noted
earlier, such studies may ultimately aid in the utilization anddevelopment of such marine plant resources.

Laminaria
In contrast to the substantial production of Laminaria in
China (Cheng 1969) and Japan (Hasegawa 1976), this species is
not cultivated commercially in North America, presumably
because there is little market for its product (e.g., kombu).
However, there is considerable interest in its ecology and productivity because of the plant's major role in nearshore
ecosystems. Much of the recent work on Laminaria in North
America and Europe (Boden 1979; Kain 1979; Luning 1979) has
direct applications to its potential mariculture and resource
management, and will be reviewed herein.
The productivity and growth strategy of L. digitata, L.
longicruris, and Agarum cribrosum were studied in St. Margaret's
Bay, Nova Scotia (Mann 1972a, b, 1973; Mann and Chapman1975). The growth of individual plants was measured periodicallyfor 2 yr by recording the distance between holes thatwere initially spaced at 10 cm intervals along the blades. As the44

ovember significantly reduced growth rates. The lower rate ofrowth was attributed to a reduction in photosynthetic areaather than a loss of storage products. An additional description
f the annual carbon budget for L. longicruris in St. Margaret's
ay was given by Hatcher et al. (1977).
The seasonal patterns of nitrate uptake in L. longicruris were
etermined by Harlin and Craigie (1976), based upon populaions from St. Margaret's Bay. Their experiments suggested that. longicruris was capable of using nitrate under winter condiions, and that it did not depend upon stored compounds for anitrogen source during periods of rapid growth, i.e., late winternd spring. Discs were punched from the thin edges of L.
longicruris plants and they were subjected to external nitrate in aseries of concentrations under different light and temperatureegimes. The optimal rate of uptake, with summer material(June and July) was 7-9 ,.MIg dry wtlh at 15 DC and approx'mately 700 fc. At 5 DC, uptake was 60070, and at 0 DC it was 40%of that at 15°C. The uptake in the dark was 60-80% of that in
he light. Algae pretreated for 12 h with antibiotics removed
nitrate from the medium at a rate of 84% of the untreatedaterial. Nitrate uptake was inhibited in proportion to the concentration of nitrate in the medium. Further studies of nitrate
uptake by L. longicruris have been reported by the same investigators (Harlin and Craigie 1978).
The growth and production of L. longicruris are interrelated
r:th nutrients, water motion, and other parameters (Gerard and!v~ann 1979). That is, measurements of internal inorganic
nitrogen (i.e., nitrite and nitrate) reserves and dry organic weight
Indicated that low concentrations of dissolved nutrients insummer and low levels of illumination in fall and winter werebore limiting to growth at exposed than sheltered sites.
The morphological differentiation of nondigitate Laminaria
populations from the North Atlantic was evaluated by Chapan (1973a, b), who found that the occurrence of long, hollow
tipes (the L. longicruris types) decreased with increasing
xposure. The differentiation of populations in relation to
xposure was clinal with overlap in the variation patterns. The
ajor taxonomic finding of the analyses was the lack ofnonoverlapping criteria by which L. agardhii and L. longicruris
ay be delimited. Subsequently, Chapman (1973b, 1974a) has
conducted heritability tests between the two morphologicaltypes within the Simplices section of Laminaria. The productionof fertile hybrids established interfertility between populations
from the extremes of the morphological range . Reciprocaltransplantation experiments between exposed and sheltered sites
produced no significant changes in stipe morphology. Quantitative genetic analysis of the degree of resemblance between
relatives indicated a high genetic component in the interpopulation differentiation . Estimates of within-population heritability
at the exposed site were low, perhaps because of intensive local
stabilizing selection which reduces genetic variability.
The inheritance of mucilage canals in Laminaria, section
Simplices, from eastern Canada has also been described by
Chapman (1974b, 1975). A matrix of heritability tests showed
complete interfertility between all mucilage canal types, fertile
F\ hybrids being produced in all cases. Quantitative genetic
analysis revealed a large environmental component in thephenotypic variance of degree of mucilage canal development.
Only plant from ova Scotia bred true with respect to mucilage
canal . Thi characteri tic was considered un uitable for taxonomic and biological specie determinations, though there \\asevidence for intraspecific genotypic differentiation in orne
ova Scotian population. Li.ining et al. (19 ) conducted cro -ing experiments in the nondigilate c mple of Lammona fr mboth sides of the Atlantic . Lammuna longl nms from\
Scotia and two forms of European I . 0 channa were ucessfully crossed. Parental and hybrid porophy1
were
cultivated in the sea near Helgoland, \\ here the> reached m
imum frond lengths of 40-120 cm. her 7 mo all e penmental
plants were nonsporogenous and were remO\ eel from the ea
The authors di cussed a vanety of obsen alion on genelJ all)fixed traits and problems of the relatiomhir bet\\een n ndigitate Laminana forms from the eastern and \\e tern Ide of
the North Atlantic Ocean.
A quantitative genetic analysis of alginate content In 1.
longicruris was conducted by Chapman and Doyle (1979), In
order to partition the plant's phenotypic \ariance Into genetic
and nongenetic components. Regres Ion anal) I of mean offspring on midparent values and artificial selection were u~ed toestimate heritability and realized heritabilit y, respectivel). Bothestimates showed that there was a low additive genetic comronent in the phenotypic variance and that a simple ma~s selectIOnprogram would produce a very small genetic response.
A summary of the experimental and numerical taxonomy of
the Larninariales was given by Chapman (1978). He de\cribeel
crossability tests, transplant expenments, standard laborator)
cultivation, and quantitative genetics of the Lammariales, muchof which is relevant to the genetic improvement of kelp andtheir ultimate mariculture. In most cases the crossability tc
showed relatively easy hybridization between taxa ha\ ing clo eorthodox taxonomic affinities. Conversely, distantly related
taxa, in a classical taxonomic sense, sho\\ed little or no
hybridization. In the same way, transplant expenment ha\edemonstrated that some closely related taxa were the phenotypic
manifestations of plastic responses to diverse habitats
The destructive grazing of kelp beds by common green urchins, Strongylocentrotus droebachiensis, and its relation hip
to lobster, Homorus american us, abundance have been reported
in Nova Scotia (Mann and Breen 1972; Breen and ;\Olann 1976a,b; Mann 1977). As in other geographical area ( orth 1971) ea
urchins have a major effect on the abundance of kelp , ~ wellas on preventing the return of kelps with contmued ea urchm
settlement. Lobster harvesting was shown to have mterrelated
effects on the abundance of kelps. For example, in one Ite In t.
Margaret's Bay, the abundance of lobsters decreased by nearh
50% in 14 yr. The lobster is a major predator of ea urchm ,andduring the past 6 yr (1970-76) the sea urchm~ ha\ e d troyed
70% of the beds of Lammaria spp. in the area. A the kelp bed
are destroyed, their high primary prodUCtlVlly I 10 t (Mann1972b), which will cau e sub lantial chang at higher trOPI
levels. The authors suggest that a management program rna be
required to increase the abundance of lob te~ In order to pro
teet the remaining kelp-bed commumUes.
The Lominaria spp from the Pa lfie Oa! t of orth menca
have also been exten i\el) tudied. For e ample, the role ofotters m structuring near hore elp communlU , In udm
Laminano spp., has been exten l\eI) tudled (Davt n 19 4Estes and Palmi ano 1974). In addition, a ummary of the elp
species and their dlStnbuuon 10 the n rthPlined. In relation (0 <x-eanographl~ ond(01969,19 o. 19-2) A detailed autecolo cal tud ofdigitate Lammar/Q pecles from[ern C nad(Druehl 1965, I - ; Druehl and H I ) n
tolerance ludiC of gametoph)1\\ ell a· [udlC~ of the ph> 1010
and [ran plant ecoloof rna ro
4'i1974, 1976; Lobban 1975, 1977, 1978b; Schmitz and Lobban
1976; Whyte and Englar 1976b, 1978; Fankboner and de Burgh
1977; Schmitz and Srivastava 1979a, b; Wheeler 1980a, b), andnutrient uptake (W. Wheeler 1976,1977; Jackson 1977a, b; Northand Wheeler 1977; P. Wheeler 1977, 1979) should be consulted, as
they also contribute substantially to our understanding of theplant's biology, management, and potential mariculture.
The southern California kelp beds, Macrocystis pyrijera,
have undergone a rapid deterioration since 1940, the kelp crop
being at an all-time low in the early 1960's (North 1971, 1976;
Branning 1976). An extensive study was conducted by Northand his associates from 1956 to 1962 to determine the cause ofthis decline (North 1971). Their studies indicated that many
fomler kelp areas had become dominated by populations ofsmall purple urchins, Strongylocentrotus purpura Ius, which
grazed seaweeds and exhibited a preference for M. pyrijera. In
addition, a decline in an important predator of the sea urchin,
the sea otter, was documented (Leighton 1962; Leighton et al.
1966). In 1962, the Kelp Habitat Improvement Project was initiated (North 1972c, 1973) in order to control urchin populations and to restore the vanishing kelp beds. The fust large-scaleurchin control experiments at Point Lema, Calif., were very
successful. However, similar experiments at other sites met withvarying success. A number of factors influenced the outcome ofa given urchin-control operation. For example, juvenile M .
pyrijera plants did not appear in abundance unless a stand of
adult plants occurred near the area cleared of urchins. Kelp
spores can only propagate the species within a few hundred feetof an established stand (Anderson and North 1966, 1967). Eventually only the urchins close to adult M. pyrijera plants were
removed, allowing juvenile plants to proliferate. Thus, urchin
control was primarily initiated around the periphery of the beds,
gradually permitting the colony to expand.
A detailed description ot the Macrocystis cultivation programs, developed during the Kelp Habitat Improvement Projectstudies, was given by North (1972b, c, 1973, 1974, 1976, 1979b).
He emphasized that a natural evolution of mariculture techniques occurred during their programs. Initially, transplant

distribution, abundance, and phenology of the kelps are interrelated with their differential tolerances to temperature andsalinity. Axenic culture techniques were perfected (Druehl andHsiao 1969) to evaluate the control of gametogenesis (Hsiao andDrueh,l1971, 1973a, b,; Hsiao 1972) by such factors as light, different culture media, nutrient content, and halogen concentration. Furthermore, the in situ development of gametophytes andyoung sporophytes of L. saccharin a was described (Hsiao andDruehl 1973c). Zoospores were labeled with fluorescent stain(calcofluor white), cultivated, and recoved (examined) at frequent intervals. The gametophytes produced gametes
throughout the year. However, the production of macroscopic
sporophytes appeared to be limited by survival of microscopic
sporophytes, particularly during the summer. Macroscopic
sporophytes were evident during the late winter and autumn.
In addition to the references summarized above, the reader
should consult the extensive review of Laminaria given by Kain
(1979). She summarized information on the taxonomy anddistribution, life history, structure, physiology, and ecology,
much of which is relevant to the future mariculture and resource
management of this plant.

Macrocystis
Macrocystis is one of the most important natural resources on
the Pacific coast of North America, for it is both a major sourceof alginic acid and a significant component of the nearshore
kelp bed community. In addition to North's (1971) extensive
monograph on the plant, several recent papers dealing with thegrowth (Neushul et al. 1972; North 1972a; Coon 1974; Coonand Wheeler 1976; Neushul et al. 1976; Jackson 1977c; Neushul
1977 In press; Neushul et al. 1977; Lobban 1978a, c; Coon 1981),
reproduction (Chi and NeushulI972; Gherardini and North 1972;
Neushul 1974, 1977; Sanbonsuga and Neushul 1978, 1979, In
press), ecology (Charters 1974; Harger et al. 1974; North 1974;
Pace 1974; Rosenthal et al. 1974; Foster 1975a, c; Harger 1976,
1977; Pearse and Hines 1977a, b; Rhoades 1977; Devinny andVolse 1978; Luning and Neushul 1978), physiology (Wheeler

Kelco's kelp-harvesting boats in a southern California kelp forest. Photograpb by A. Mathieson.

46

techniques were attempted with large Macrocystis plants (North1964), particularly in areas where the giant kelp had vanishedcompletely. Such transplants were very laborious and often
frustrating, for many plants did not survive. In some cases, thetransplants were too small to achieve ecological stability, due toheavy urchin and fish grazing. North (1976, 1979b) summarized
his transplanting techniques for macroscopic plants, including
descriptions of the anchoring systems used, transporting procedures, and optimum plant size for viable transplants. He indicated that in one joint effort with Kelco Corporation (a kelpharvesting company in San Diego, Calif.), approximately 20,000
juvenile kelp plants were transplanted and installed torepopulate one kelp bed in La Jolla.
Ultimately, mass culture techniques were developed which incorporated the use of microscopic sporophytes as seed fornatural reforestation (North 1972b, 1974, 1976). One very successful technique used in the program involved the attachmentof spores on strips of polyethelene fIlm 8.5 m long, folded inpleats so as to fit in a refrigerated aquarium with a dense inoculum of spores. After the spores had settled on the film forseveral days and developed as gametophytes, the fIlm wastransferred to a sloping tray with a swiftly flowing seawater
source. Juvenile sporophytes were produced after only 1-2 wk
because of enhanced growth due to the vigorous water motion.
The external surfaces of the sporophytes are sticky. Accordingly, sporophytes can be scraped off the polyethelene strips andimmediately dispersed in the sea, or if necessary, held in culture
prior to dispersal if aerated and kept in suspension. North (1973,1976) estimated an average survival rate of 11100,000 with thepresent techniques. Consequently culturing techniques weredeveloped to produce billions of sporophytes very inexpensively.
He emphasized that the seeding techniques should be
economically feasible for use in the United States, with its highlabor costs.
A number of factors were shown to influence the sporophyte
dispersal operations, including seasonal variability of the seed,turbidity, sedimentation of rocky surfaces, and the age andamount of seed'used (North 1976). The summer and fall were
the poorest seasons for dispersing sporophytes. Unfortunately aconsiderable length of time (3-4 mo) was required before thesuccess of a seeding experiment could be evaluated. An additional 6-9 mo must elapse before a juvenile kelp develops sufficient canopy to be visible.
A variety of other transplant techniques were attempted(North 1972b, 1973, 1976), including the growing of juvenile
kelps on plastic (PVC) rings (\1.1-12 inches in diameter), which
could be attached to a rocky bottom by epoxy resin. The
method involved considerable labor so it was used only in
special circumstances. For example, such techniques were
employed to select for heat-tolerant strains. Such strains exist
primarily in southerly regions. They showed good potential astransplants to areas where heated discharges were produced
from coastal generating stations (North 1971), thus creating kelp
beds with considerable immunity to thermal effects.
Devinny and Leventhal (1979) have also outlined several new
methods for the mass culture and dispersal of juvenile M.
pyrifera sporophytes. Basically, they have attempted to reduce
costs for laboratory space, refrigeration, and water flitration,i.e., as compared to those previously developed by North(1976). In addition, diatom contamination was successfully controlled by washing cultures with turbulent seawater.
Other summaries of the kelp restoration program in southern
California have been given (Wilson et al. 1977, 1979). The latter

investigators suggested that although techniques have evolved tothe point where it is possible to strongly influence competingvegetation and sea urchin populations, the ultimate success of akelp restoration effort will be the identification, protection, andmaintenance of a healthy environment.
As noted in the section dealing with Marine Biomass, there
has been a rapid enhancement of techniques for the domestication of M. pyrifera (NeushuI1977, In press). Even so, there areseveral priority areas necessary for the mariculture of the giant
kelp, including selective breeding of fast-growing strains, thestorage of germ plasm of genetically identical progeny, and adetailed understanding of the recruitment and loss of juvenilestages. With respect to the first two priorities, a greenhouse andtank complex at the University of California at Santa Barbara
have been perfected (Neushul 1977, In press; Neushul et al.
1977; Charters and Neushul 1979) where it is possible to raisekelp sporophytes to reproductive maturity and to make controlled crosses. With such a system various crosses can be initiatedand their progeny raised to verify and evaluate the heritability ofgrowth characteristics. Intergeneric hybrids have been established between M . pyrifera and Nereocystis leutkeana and between
M. pyrifera and Pelagophycus porra (Sanbonsuga and Neushul
1978, 1979, In press). In each case the hybrids were very different from either parental types. Neushul (1977, In press) has
further suggested that while the implications of intergenerichybridization are not yet fully appreciated, the ability to produce these plants implies that Macrocystis itself, when known

Seaweed cultivation facill!) at Unher;it) of Oilifom... iOla Barban.
sho"ing • close-up of kelp culti,ation in • long. /'Klangular t.n~ ..ith •"water-broom" aenltion unit. Pbotognlpb b) 4. \lathle>On47

Cultivation of Eucheuma uncinatum at University of California, Santa Barbara. Photograph b} A. Mathieson.

of nitrate-N, as well as ammonia and phosphate, varied withlight intensity and nutrient concentration. P . Wheeler (1977)
summarized further infonnation on ammonia-N uptake rates inM. pyrifera.
The growth of M. pyrifera is limited by the availability of certain nucronutrients (North 1977c). Renewal of nutrients otherthan nitrogen and phosphorous probably depends on runoff
and local regeneration from decomposition processes. According
to North, M. pyrifera increased its nitrogen content whentransferred to nitrogen-edriched media, which suggested astorage function. Thus, lertilizing strategies should consider
these characteristics, and certain classes of limiting
micro nutrients might best be dispersed continuously whileothers applied only as needed. North further emphasized thatcaution must be used when delineating the nutritional quality ofseawater by inorganic nitrogen, for micronutrients may belimiting. That is, experiments have shown that growth rates ofjuvenile Macrocystis sporophytes in seawater are greatly
stimulated by proper nutrient enrichment (North 1977a, b). Fertilizing programs should foster the production of commercial
kelp farms (North and Wheeler 1977).
Further descriptions of the growth of M. pyrifera have been
given by Jackson and North (1973), Bryce (1977), Flowers andBryce (1977), Jackson (1977c, 1980), Neushul (1977, In
press), and Hart et al. (1979) in the section dealing with Marine
Biomass. A variety of biological, technical, and cultivation infonnation is summarized by the above investigators, in orderto evaluate the production potentials of methane from kelp
biomass.
A Sea Grant project by J. Pearse at the University of California, Santa Cruz, evaluated the population dynamics and interactions within the giant kelp forests off the central California
coast. One of the major differences between the central andsouthern California kelp beds, other than hydrographic variations, is the presence of sea otters in some of the central California forests (e.g., the Hopkin Marine Life Refuge) and their imminent establishment in others (e.g., Santa Cruz). Basically,
Pearse and associates studied the productivity of the giant kelp
(Towle and Pearse 1973), as well as the population dynamics ofvarieties are crossed and the progeny grown, will be amenable togenetic marupulation. In addition, it has been possible, using appropriate culture conditions, to maintain Macrocystis
gametophytes for prolonged periods in culture vials, thereby
providing a means of stockpiling genetic strains as a gene bank.
Several fouling-plate studies have been conducted to evaluate
the in situ growth, recruitment, and survival of seaweeds, including M. pyrifera (Harger et al. 1974; Neushul 1974, 1977;
Harger 1976; Neushul et al. 1976). The fouling plates werehorizontally positioned to concrete parking bumpers and attachedon the ocean floor ( - 12 m); they were periodically taken fromthe sea, subjected to nondestructive microscopic survey in thelaboratory, and then replaced in the ocean in order to quantify
the growth and reproduction of seaweed populations. Neushul
et al. (1972) made direct microscopic observations of algal spores
under natural conditions by taking a microscope into the sea
that was equipped for incident light and high magnification.
The techniques of outplanting and microscopic examination in
the laboratory and field should provide detailed understandingof the in situ biology of seaweeds such as M. pyrifera.
The interrelations of nutrients and productivity with M.
pyrifera were determined by Jackson (1977a, b). The mostlimiting condition for M. pyrifera was the low concentration ofdissolved nutrients, especially nitrogenous substances.
Translocation of nitrogenous compounds by the kelps from
depths where concentrations were higher compensated for thislimitation.
Other accounts of nutrient uptake kinetics in M. pyrifera havealso been summarized. Haines and Wheeler (1977) showed thatM. pyrifera exhibited saturable kinetics for nitrate-N. Ammonia
uptake showed saturable kinetics at concentrations below 22
pM; at higher concentrations the uptake increased linearly.
Nitrate and ammonia were taken up simultaneously; thepresence of one fonn had no effect on the other. According to
W. Wheeler (1976, 1977), the photosynthetic rates ofMacrocystis increased in situ as the rate of water flow increased
from 0 to 6 cm/s. The development of a turbulent boundary
layer adjacent to the blade at such low water velocities wasenhanced by the corrugate morphology of the blade. The uptake
48

cultures were included as a final phase in the system (Haines
1975; Haines and Monahan 1976; Roels et al, 1976a, b; Hainesand Wheeler 1977), In order to recover some of the nitrogen
losses from the system and to reduce a potential environmental
hazard, the carrageenan-producing red alga Hypnea musciformis was grown very successfully in the effluent from shellfi htanks, Hypnea grew five times faster in the shellfish tank effluent than in unaltered deep seawater and about three times
faster than in surface seawater (Roels et al, 1976a). The seaweedgrown in the effluent doubled its weight every 60 h (Haines
1975), The average ammonia-N utilized by the seaweed was
511110, varying between 41 % in one study (Roels et al, 1976a) and
61 % efficiency in another study (Roels et al. 1976b), Higher efficiency of arnmonia-N utilization would be possible if a slower
turnover volume were employed, i,e., < 95/d, A variety of otherdetails regarding the growth of H. musciformis in the artificial
upwelling system at St. Croix (Haines 1975) were given in thesection dealing with Hypnea,
Economic seaweeds were employed in the deep-water upwelling mariculture system, in order to reduce its costs (Roels et al,
1976a) , For example, the potential yield of H. musciformis
alone would be 165 tons dry wt/ha per yr; at $650/ton this would
represent a yield of $107,250/ha per yr.

important invertebrate grazers and their predators (Pearse andHines 1977a, b). A detailed description of the densities, sizes,
and distribution of abalones and sea urchins , as well as thespatial distribution of grazing 'snails, was summarized (Lowry
and Pearse 1973; Lowry et al, 1974), Pearse's studies have provided a unique opportunity to document changes in kelp bed
communities (e,g" Santa Cruz) after the reestablishment of animportant carnivore-the sea otter (Pearse and Hines 1977a).
Such studies will ultimately provide the information needed forpredictive manipulation of the complex interactions within kelp
forests. For example, the giant kelp provides the major trophic
base, particularly in the form of debris (drift) and detritus, formany animals within the kelp forest, including economically important species such as abalone and crabs,
The potential use of NASA-LANDSAT data to make annual
evaluations of the sea otter-urchin-kelp relationship was outlined
by Rhoades (1977), He suggested that an annual census of thekelp canopy in California could be prepared to evaluate thestability and resilience of all kelp beds from Point Sur to Diablo
Canyon.
A variety of studies were recently initiated in British Columbia, Canada, in order to enhance the management of M. integrifolia (Druehl 1977, 1978, 1979; Druehl et al . 1977). The
species is restricted to regions having little seasonal variation oftemperature and salinity or sites where the lower salinities occur
during the winter (Druehl 1978), A provision of substrata adequate for the giant kelp but inadequate for herbivorous invertebrates , such as urchins and turbon snails, allows theestablishment of new beds, The distribution of photosynthetically fixed carbon, respiratory requirements, exuded
dissolved carbon, and detritus were determined (Druehl 1978),As a result, the standing crop of M, integrifolia was circumscribedin terms of total biomass, pigments, carbon, nitrogen, andalginic acid, Major invertebrate and vertebrate inhabitants of
the kelp beds have also been defined, as well as the food chainsoriginating from M, integrifolia (Druehl et aI. 1977). Druehl(1979) emphasized that an understanding of the total ecology of
giant kelp beds in British Columbia was necessary for continued
management of this valuable natural resource,

Marine Biomass
The Office of Technology Assessment of the U.S. Federal
Government conducted a feasibility study, "Energy From
Marine Biological Processes," at the request of the Senate Committee on Commerce (Committee on Commerce, Science, andTransportation 1980). The study, which was also referred to as the"Energy From Marine Biomass," addressed a variety of topics asfollows: 1) The potentials for specific seaweeds as biomass producers; 2) the status of the technology of ocean farming; 3) the
status of marine plant culture research; and 4) regulatory, environmental, and social impacts. In addition to the above investigation, several other studies have also evaluated the potentials
of seaweeds for marine biomass production and/ or the conversion of marine biomass to methane (Jackson and North 1973;
Leese 1976; Bryce 1977; Flowers and Bryce 1977; Goldman and
Ryther 1977; Krauss 1977; Mitsui et al. 1977; Neushul 1977, In
press; Anonymous I978c; Hruby 1978; Michanek 1978; North
1978, 1979b; Hart et al, 1979; Committee on Commerce, Science,
and Transportation 1980; Doty In press b; Jackson 1980; Rao et
al, 1980; Hanisak 1981), Some of the major conclUSIOns ofthese investigations are given in the subsequent paragraphs of thissection.
An extensive review of the problems and potentials of marine
biomass production through seaweed aquaculture was given byJackson (1980) and Jackson and North (1973). The) emphasized the need for more basic physiological and ecological information in designing seaweed aquaculture systems, panicularly modeling evaluations. For example, little is known about theinteractions of high temperatures and low nutrient concentrations, two factors that could be separately controlled in an
aquaculture facility. A simple model was presented whIch suggested that yields of the giant kelp, Macrocystis pyrtjera, could
be increased 50% by increasing water temperatures 10
Observations about the size and type of culture facilities werealso reviewed by Jackson and orth (1973) . The) uggested thata closed system would require an exchange of water atleasl once
per day, as slower exchange rates .... ould lead to carbon depletion by seaweeds and unwanted ph)toplankton accumulation.

SELECTED MARICULTURE PROGRAMSThis section describes the growth and potential benefits ofseaweed populations in three mariculture programs, That is, themariculture of several economically important seaweeds is
described at the artificial upwelling system in St. Croix, U.S.
Virgin Islands, as well as in association with the Marine Biomass
Program and several polyculture waste-recycling projects.

Artificial Upwelling Systems
The improvement of seaweed growth by supplying nutrientrich deep water from an artificial upwelling system should becited as a good technological application in mariculture. A
detailed description of an artificial upwelling site at St. Croix,
U.S, Virgin Islands, and its application to mariculture was summarized (Roels et al. 1970; Roels et al. 1971; Roels 1975;
Sunderline et al, 1975; Sunderline 1976), Basically, nutrient-rich
deep water (870 m) was pumped into ponds (45,000 liters) where
phytoplankton were grown as food for ftlter-feeding shellfish.
The deep water employed in the mariculture system was free ofpollutants, parasites, diseases, and predators, In addition to theprimary unialgal culture system described above, seaweed49

method for \ccding M . p)'rtjera on natural ub\tfata in the~,North (1976) has ulrcudy IIJ.lde u)flslderahle progr
In Ihi~ area .
An evaluul10n of the recruitment and 10 of juvc rllle la es I an
importanl con ideration In future rn na (.'ment and Jluvation.
Thus, Neu hul (1977, In pr s) uggcsted that the ellpcrimental
manipulation of fouling-plate \X)PUI,llIon of M . [lyrijero, awell as other plant, would 1I0w Iderlltfic. ti n :tnd ev rIlual
control ot the "envlfOllm::nt I turn til .. postul<ttt.'d Y 'orth
to contri ute to the high mortality of )'oun por lphytes 10 thesea.
de cripllon of the research nd development pr
am toa ~. the economiC fe Ibillty of methane production from M .
pyrifera \\'ummarllcd by Bryce (1977). Palo pi nt ev luaU n were J\cn in vunety of area : I) ontro11OO cultivation in
10 of rav. matenal tothe open 0 can; 2) pretreatment pr
enhance anuerobi digestibility, J I lion of peclfi ana roblc
microorgani m for kelp dcgradall n; 4) opllmll4l1lon of thefor production of commerClal quananaerobl dl lion pr
title of meth.lOe ga ; ancl S1 de el pment of a y tern model for
economi aud techmcal
u tI n of the manne farm con·
cept .
A fermentation pr
methane from \fpyrtfera v. descnbed (Hhlch reduced the 10hlbitory or to I level ofter content, and tncrea 00 bul den It} . Pretreatment, an the presence of aO"
was follov.ed by compr Jon to rc:mo\e 75 0 of the mOl tureand 70 11'0 of the a h JmllaIl} present, htle ret 1010 fO~O of thevolatile ohd for ermentauon
The potenll I 0 algae for entrappm
olarphoto }nth I were umm'nzed by Dot} (In prb). Hegest d that the ph t yntheu entrapment potmllal
difli ult bccau e of the scale required to obtain th\\ d and Benemannmou amount of ener needed
hale uggested that the uuluallon of all the
e \\ te tn thefertthzer \\ ould prol1de < I ~o of thenlled ' tates
energy need . fo meet th e need (Hall I If) ome ,9km' of effiCIent algal farm I\ould be requlfed A ordingJy.
DOly (In pre b) uggested that considera Ie ph}cologrcal manpower would be nec aT) to me t the orldwidt: energ) needof the future.
A I'ariet) of other Infomlution on marine bioma \a ummarized at a recent conference entitled "The EconomiL andEngineering of Large-Scale Igae Biom~ ) tem ." held at theMas achusetts Institute of Technology during JanuaT)' 19 .
The conference reviewed the tate 01 the an on vadou marine
biomass systems (Anonymous 197 b).
synopsi of theworkshop on fuels from manne bioma pointed out that I,
there was potential error in extrapolating today' mall y terns
to the prOjected 100 mi' farm becau e of the relatively primItivestate of the an in mass culture of algae; 2) the de igning,building, and operating of a large algal biomass y tem olely
for the energy product would be prohibimely expen ive basedon current knowledge of mass algal culture and e.xtrapolations
from this knowledge; also, such a system would pro\oke considerable environmental concern, because of competing uses forland and water; 3) for very large systems, pro\iding the required
nutrients would be the major engineering and cost problem; 4)basic research in genetic engineering would be needed to obtain
additional valuable end products, such as feedstocks and protein, in addition to energy, as economic justification of massculture; 5) small, I-mi' systems using sewage sludge as nutrients
may merit further study; 6) the advantages of disposing ofundesirable sewage sludge may, in site-specific cases, make

Carbon concentrations can be regulated by bubbling carbon
dioxide through the system, but it would be prohibitively expen
sive. Oceanic culture facilities would have to be I km or more in
length, jf the nutrients were to be efficiently used by the desired
seaweed. However, if an oceanic system becomes too large, its
flushing requiremen ts would be similar to those of a closed
system.
Additional information regarding the selection of the giant
kelp, M. pyn!era, for possible cultivation on offshore mannefarms has been summarized (Jackson and North 1973). The major advantages of M, pyrijera, as abstracted from Hower andBryce (1977), are as follows: I) The giant kelp is ea ily cultured
and has a relatively simple hfe cycle; 2) AI. pyn!era is sufficiently
tough to resist rough water conditions and han'esting strc e~; 3)
the size of the adult plant IS substantial. approximately 60 m.
and it requir s a minimum of supporting struc( ure tor it
cultivation; 4) /\1. pvrijera make maximum utilization ofsunlight at the surface and is therefore relatiyely effiCient at capturing solar energy; 5) the morphology ot the giant kelp ullo\\s a
portion. at the surface, to be han'ested. lea\ing the remamder toreplace lost tIssue, thus replanting after harvest is nnt rCQuired;6) the flotation mechanism of .'1-1 pryifera facilitate mechanicalhanesting . Three critical con ideration' involved in ulti\utlonof M. p~'fijera are growth substrata. nutrient suppl~, andhan'esting
The Em:rgy From Marine Biomass program ha as it 01 crallgoal the de termination of the technical and economic feasibllttyof a commercial-sized system for the production of methanefrom kelp biomass (Anonymous I977c; Flower and Bf) e1977), MacrocystlS could be cultivated on artificial ub trata inthe open ocean and harvested and converted b} biological or
physical/ chemical processes into substitute natural gas. Rc idual
by-products of commercial value would be separated, further
processed, and used for feed and fertilizer.
A quarter-acre module (QAM) marine farm has been deSigned to demonstrate the feasibility of growmg and han csting
giant kelp in the open sea (Flowers and Bryce 1977). The marine
farm would be moored in a minimum of 500 ft of water, approximately 5 mi offshore from Corona Del Mar, Calif. A
substratum structure, consisting of a tension grid system, wasselected for the marine farm. This system is a net like unit hal ing
discrete, flexible cable members which are capable of supporting
loads by axial tension. Flowers and Bryce summarized two
general approaches for the conversion of Macrocystis tomethane: Anaerobic digestion and hydrogasification. Sinceanaerobic techniques are fairly well defined, while processes forhydrogasification of biomass are just now being investigated,
the program emphasis is placed on biological conversion. Further details regarding methane generation and the different
phases of the Energy From Marine Biomass program are summarized. Prelimianary approximations indicated that the costsof methane from Macrocystis biomass would be in the samerange as gas produced by coal gasification.
The domestication of M. pyrifera as a marine plant biomass
producer was discussed by Neushul (1977, In press). He emphasized that preliminary research in the use of seaweeds as collectors of energy is underway (Jackson and North 1973), andwith improved techniques, progress in domestication will be
rapid. Of particular importance will be the long-term storage ofgerm plasm or single-cell isolates (gametophytes) of genetically
identical progeny, similar to that described by Saga et aI. (1978)
for clones of single isolated cells of Laminaria. A number ofother problems exist, including the necessity to improve the50energy production commercially viable. An opportunity briefsummarizing many of the above described economic andengineering considerations associated with algal biomass energy
systems was recently publisheo (Anonymous 1978c),

marized by Ryther (1975, 1977). Successful cultures ofunicellular algae and seaweeds were sustained for several
months with only minor problems. Algal yields and nitrogenremoval capacity varied seasonally by threefold to fourfold andappeared to be primarily controlled by solar radiation. The
combined unicellular algae-shellfish-seaweed system wascapable of completely removing nitrogen from wastewater effluent within an area of 48 acres. Ryther estimated an annual
production of 183 tons of oyster meat (38,000 bu of wholeoysters), 3,350 tons (wet weight) of seaweeds, and undetermined
quantities of flounder and/or lobsters per treated effluent of
10,000 people. The seaweed system alone was capable of complete nitrogen removal in an area of 60 acres with an annual production of 16,300 tons wet weight. The above figures were for
year-round operations such as would be possible in tropical tosemitropical climat("s. Operation of the system in temperate
climates would be possible within the same areas but only on aseasonal basis (approximately 6 mo), with one-half the above
yields.
The removal efficiencies (i.e., stripping potentials) and algal
bioassay procedures employed for inorganic nitrogen removal in
a combined tertiary treatment-marine aquaculture system at theEnvironmental Systems Laboratory have been described(Goldman, Tenore, Ryther, and Corwin 1974; Goldman,
Tenore, and Stanley 1974). A prototype facility consisting ofgrowth containers for marine algae, oysters, and seaweed, joined in a series, was fed secondarily treated wastewater diluted 1:4
with seawater for 11 wk. During this time, 95070 of the intluent
inorganic nitrogen was removed by algal assimilation. The
oysters in turn removed 85 % of the algae, but regenerated
16-18% of the original nitrogen contents of the algal cells as
soluble ammonia. All of the regenerated nitrogen was removedby the seaweed system so that the total inorganic nitrogen
removal efficiency was 95%. Phosphorus removal, on the other
hand, was less efficient, as only 45-60% was removed. The algal
bioassay studies (Goldman, Tenore, and Stanley 1974)
demonstrated that nitrogen removal was necessary to prevent increased algal growth of coastal marine waters receiving

Polyculture
Several recent experiments involving the growth of unattached economic seaweeds have been conducted in the outdoor massculture facilities (tanks, raceways, and ponds) of the Environmental Systems Laboratory of Woods Hole, Mass., and of
the Harbor Branch Laboratories in Fort Pierce, Fla. (Ryther et
aI. 1972; Anonymous 1973a; Fralick 1974; Goldman, Tenore,
Ryther, and Corwin 1974; Goldman et aI. 1974; Prince 1974;
Ryther 1975, 1976, 1977; Ryther et aI. 1975; DeBoer, D'Elia,
and Lapointe 1976; DeBoer and Lapointe 1976; DeBoer, Lapointe, and D'Elia 1976; DeBoer, Ryther, and Lapointe 1976;
D'Elia et aI . 1976; Lapointe and DeBoer 1976; Lapointe et aI.
1976; DeBoer et aI . 1977; DeBoer and Ryther 1977; Nicotri
1977; Hanisak et aI . 1979; Ryther et aI. 1979; Stenberg 1979).
Basically the studies incorporated seaweeds as well as
phytoplankton in a waste-recycling marine polyculture system,
which also included fish and shellfish (Ryther et aI. 1972; Ryther
1975, 1976, 1977; DeBoer, Ryther, and Lapointe 1976; Fralickand Ryther 1976). Such a system has the potential of removing
inorganic nutrients from treated sewage effluent, prior to its
discharge into the environment, and recycling these nutrients into commercially valuable products. The basic concept of thesystem is to involve three or more cultured organisms. For example, unicellular marine algae can be grown in ponds with acontinuous flow of mixtures of seawater and secondarily treatedsewage effluent. The unicellular algae are then fed to shellfish inorder to remove nutrients from the wastewater. Finally the effluent from the shellfish cultures passes through tanks containing seaweeds, which remove the dissolved nutrients regenerated
by the animal cultures before the final effluent is discharged.
A preliminary account of the pilot plant waste-recycling
marine aquaculture system at Woods Hole, Mass., was sum-

Seaweed culture tanks at the Environmental Systems Laborntory, Woods Hole Oceanogrnphic Institute, Woods
Hole, Mass. Photogrnph by D. Cheney.

51

production during both pring (U-4J g dry wt/m 2 per d)ummer (20-36 g dry wtlm l per d). Inc productIon rat 0
tikvahtae were variable, incrca Ing from 4 g dry "-l/ml per
Apnl to 43 g dry ~t/m' per d in carly June and then deeragain during late ummer to 7-1 g dry wtlm' per d. The
duct ion rates of both SpecIes declined dunng fall and~inter . By mid-December the production rate lor I. tlkva
had fallen to lero, and the plant detenorated; '. hOlI ymained viable with a productIon rate of 6 g dry y.l/m' pe
Overall the groen onal 'han 10 Ilroduction ~ere correl
with water temperature nd mcident I r r dmtion. Howfrom I June to 1 0 tober, hen .... ater temperatures andradIation ~ere presumabl) optimal lor seaweed ge(DeBoer, I apolOte, and D'E.ha 1976), production hoy.ed
correlation WIth these vanable. Lo .... produ tion rat .... ere
probably due to the hI h aweed densitl in the raa)high as 12.3 and 10.2 k fr h ....1 ml for ' bade I an
likvahlae, respectIvely
The mean annual dr) weIght production for} . baIleyl v.
g/m' per d (63 tlha per T) \crsu <) glm l per d (33 tlha
for G. tthahlae (DeBoer et I. 197 ).
rates may be hithan tho e of a commer 1al cnterpn bealU e the ra CY.amaintained at elevated temperatures for approximately 6
the year. Annual production rat based on a 5.5-mo geo
eason ~hen the ra CY.ay y.ere not heated ( Ma)-~O 01975) y.cre 46 tlha per )T and 2 tlha per)7 for N. batle},(,. lIhahwe, r pecu\el) Cherall, the Y1dds obtained mtud}, y.ere Iml!a! to tho reponed for Gracllarla and Hy
muscijormLS m Flonda (Lap inte and DeBoer 19 6). In
mall- ale, hon-durauon c pcnments (Hain
1975; De
Lapointe, and D'Elta 1976; ruh 19, ,Waaland I ,6,
production rate of cultured eay.eed .... ere r orded
tv.
to threefold higher, hoy.e\er, th e yield .... ere not ubstantb} large- cale, long-term produ tlon tudi .
everal operational onslderation for impro\ed ea
marlculture ha\e been re\ley.ed (DeBoer and Ryther 1977).
con ideration i the maintenance of ea .... eed bioma at
sitie that provide rna '1mum yield per unit area. For e .
the density of G. lIhahloe and \'. baile) I hould be maintat 1,800-2,
g/m 1 dunng the y.1nter, 2, ()(}..4,5 gl m'dlate spring and ummer, and at intermediate den itie durin
remainder of the year. The optimum depth of the culture .
is another variable that i best determined empiricallypenence with \ arious type of culture endo ures and mod
circulation showed that the optimum depth for the t~o s
60-110 cm. Another critical operating parameter i the 0
tration of nutrients neces ary to u tain a maximum
rate. For example, ammonia-~ was limiting below O.
another study, DeBoer, Lapomte, and D'Elia (1976) detethat the half-saturation constants for gro~lh of G. tih'and N. baileyi were approximately 0.5 /lffi ammonianitrate-No Since such constants were very low when compphytoplankton, they indicated that seaweeds utilized ve
concentrations of inorganic nitrogen. Further informati
the nutrient uptake kinetics of N. bailevi was summariz
D'Elia, DeBoer, and Ryther (1976).
.
The impact of grazing by herbivorous crustaceans
phithoe and Idotea) on mass cultures of G. tikvahiae
baileyi raised in the Environmental System Laboratory
Hole) was summarized by Nicotri (1977). Grazing rates
estimated in the laboratory and approximate crustace
sities determined in the raceways. Grazing losses were genvery small in comparison to algal productivity, but they.

wastewater discharges. Assays of effluent from the seaweed
system showed that the removal of regenerated nitrogen lowered
the potential growth to that of natural seawater.
The outdoor mass culture of GraCilaria and Hypnea
ml4cijormis in the third stage of a nutrient removal-aquaculture
system at Fort Pierce, Fla., was reported by Lapointe et a!.
(1976). Mixtures of secondary-treated wastewater and seawater
constituted the initial nutrient source to the system, which included microscopIc algae and oysters The Hypnea cultun:
could not be maintained dunng the summer when tanktemperatures reached 30°C, but Gracilaria grew dunng the entire year. On the basis of periodIC harve ting, yields were kept
consistently between 12 and 17 g dry wt/m 2 per d for both
species. Such yields were as high as those achieved for masscultures of microscopIc marine algae and fa t-growlng commercia! crops such as sugar and rice.
Tl e nutrient assimilation and growth of Juvenile Chondm
crispus plants in mixtures of seawater and secondary se~age
treatment effluents y.ere summarized by Prince (1974) O\erall,
the growth of C. crispus was similar whether nitrogen Y.as provided by ammonia-N or nitrate-No Optimal gro~th of ju\enile
C. crispus plants occurred in a sewage-sea~ater mIxture upon
addition of trace elements. The plant's growth In a 10070 ey.ageseawater mixture was not inhibited, probabl} because the ammonium levels were approximately one-half thos\: of typicalsewage (compare Dunstan and 1enzel 1971).

n.ese

Three additional nutritional studies of red algae lDeBoer etal. 1978; D'Elia and DeBoer 1978; DeBoer 1979) have beensummarized, based upon culture experiment at the Environmental Systems Laboratory of the Woods Hold
Oceanographic Institution. DeBoer (1979) and DeBoer et at.
(1978) evaluated the growth rates of G. tikvahiae (de ignated as
G. joliijera) and Neoagardhiella baileyi as a function of mtrogensource and concentration. Gro~lh rates appeared to follow
saturation-type nutrient uptake kinetics for plants recei\;ng ammonium, nitrate, urea, or sewage effluent enrichments. Ammonium enrichment produced higher growth rates than nitrateor sewage enrichment The lowest groy.lh rates occurred inunenriched seawater or urea. Half-saturation constants (K) for
growth were approximately 0.2-0.4 ~t N for all nitrogenenrichments examined. D'Elia and DeBoer (1978) evaluated the
kinetics of ammonium and nitrate uptake with the same two redalgae. Nitrate-N was taken up In a rate-saturating fashion
described by the Michaelis-Menten equation. Ammoniumuptake was multicomponent, with a saturable component being
accompanied by a high K component showing no evidence of
saturation. Nitrogen-starved plants showed higher transient
rates of ammonium-N uptake at a given concentration than
nonstarved plants. Ammonium-N was preferred over nitrate-N,
even with plants preconditioned in nitrate-N as a sole nitrogensource. Nitrate-N uptake was suppressed at 5 ~ (NH. +), but
simultaneous uptake occurred at unsuppressed rates at lower
concentrations. The potentials for nitrogen accumulation weregreater via NH. + than NO, - uptake.
The production yields of G. tikvahiae and N. baileyi, whichwere grown in a waste-recycling raceway system at Woods Hole,were summarized by DeBoer and Ryther (1977). As noted in an
earlier section dealing with Gracilaria, the plant is an important
agarophyte. Neoagardhiella baileyi contains iota carrageenan(DeBoer, D'Elia, and Lapointe 1976; DeBoer, Ryther, and Lapointe 1976), and although not yet harvested, it has potentialcommercial value. Noeagardhiella baileyi had very high rates of52

ignili unl when eru I call popul II n
ere
und nt andI al produt:IIVlly wa 1m...
1 he elfect~ ot ewage cf fluent on the TO Ilh of U/\a /a fUcan a re irculalHl
ilw:tter y tem were rep ned b) GUI t andHumm (1976). Plant 10 ,10, alld 15f:'o effluent rey, pr
rc I\'ely fa tcr than control: ull plant gr "n In emu nt c In 11 e hibited highcr nitrogt:n content than Ct !ltrol
pcrimenls, ammOl1la"a rapidl) reduced to a loy, le.. el or lero,
while phosphate was not taken up morc rapIdly than It y,ac encrated. A arc (I9HO) ha; al () c,a1uatt:d thc po Iblliu oftili/ing animal wa\tt: a a nitrogcn ource lor the red aI
Gract/aria flkvafllae and '(!oagurdll/clla ballen. Both heep andabbll wa\tc~ wcrc good nutrient ourfor the eeonorl1l II)
Important \t:itwct:d~.
The nitrate uptake of Enteromorpha mfe.."mulLs and L. Ilfiza
wit\ evaluated to determine their potcntlal u<,ef ulne in 10 eel
polYlulture and tertiary \t:wage trealmcnt \ystem~ (Harlin 197 ).
Both ,pccies exhibited a rapid uptake of nitrate, particularl) at
high temperature\ (15 ' ) and illummation (ca. 1,00 JLW/elll
per \). The optimal nitrate uptake for !:.nferumorpha pp. diller
from that of ChondnLs CnsplL5, a red alga it frequcntly con
tanlmates m Lulture (Lnnght 1979) . Thu , Harlin (197 ) uggL'Stcd
that varying nutrient uptake effiCiencies ~hould b con ideredwhen deSigning feeding \trategies for aquaculture sy~tem lor e ample, nitrate-rich sea\vater lllight be pulsed through tanb of!:."teromorpha spp. during the day. while night feeding of
cmpus culture tanks would capllalile upon the dJfTeren~c If1
nitrate uptake strategies between the two algae. thereb) minlOlIZmg uptake by the nuis3m:e seawe d.
A recent ea rant study at the niversity of Rhode hland(Harlin 1976; Harlin et aI. 1979) has evaluated th potenllal
manculture of macroalgae In ~ilo-type fish culture faulitic . 1 heopomal growth of G. flh'ahlGe in large tanb containing
The seaw~'(]s were u ~'(] tosalmollld fish \\a.\ dcscribl>d.
rcmove IlItrogcnou\ waste~ from the culture syst rm and to produce a commerCially \ aluable polysaccharide (agar) as abproduct to an existing indu5trJ (t\1eade 1976) . The plant appeared to preferentially remo\ e ammonia- . as mtratc- \\only removed after ammonia disappeared. In addition.
ammoma\\as only remQ\ed \\hen there "as nevv gro\\th on
the thallm. I.C • \\ hen ne\\ internal pool of ammoma-. vvere~ t,lhli,hL'(]. 1 he uptal-e ratc (}\m) of anlOlllma·
\\.1
hght
dcpcndent and influenced h) pretreatment condition 1 unher
H1formatlon on the ammonia uptake char (ten til. of G
ri/"\'Ohiac \\Jth rc. peet to light inten,1t . temperature. nd
ammoma- pretreatment In cis \\ er~ ummanled b) Ventura
and Harlin (1976) (also n {ll11notc II).

lMM

2

3.

4

fnt/{/cum6.

at Halifax Laboratory, R~arch and Development Directorate, F~henes
and Manne Service, EnVIronment Canada, Halifax, .S., Jan . 3, 1975,
18 p. (Mlmeogr )I977a. Mariculture<ultlvatmg the crops of the ~a , (In [,ng!. and Jr.J
Sci Dimcos . 9(4):8· I J.
1977b . New tank farm hopes to prOVide a1ternallve 50urce for In h mlXl.
Food Eng. 49(3) : 102 103.
I 977c . Kelp farm planned for methane project Ocean Jnd 12(IO)67~.
I977d . Closed<ycle aquaculture ,'I-lalS. Imt fechnol Sea (,rant Pro>
gram, Mar Ind . Advls. Serv., opportunity brief n. Rep . 77·15, 30 p.
1978a. Aquaculture 10 the UOIted StJltes, con ratnU and opportunllJes.
A report of the Committee on Aquaculture. Board on Agnculture andRenewable Resources . Comm Ion on Natural Resources, atlona! Resource Council
Natl Acad. Sci" Wa\h., D .C , 123 p.
1M.
1978b. Manne blomalS Investlgaled as a1temall'e energy lOurce.
Technol Soc., New Eng!. 5«1., New I 3:3.
197&. The econorrucs and engmeenng of large-scaJc algae blomac energy
systems. Ma s. In t. Technol.
a Grant Program, Mar . Ind . Ad,,,.
Serv, Rep 78-11,29 p.
ASARE, S.
1979a. Nitrogen and phycocolJold relations 10 Graclfano IIkvahwe and
Neoaflardh,el/a ba,ll!)!l (Rhodophyceae) J Phytol. I 5(Suppl.): I 8.
I 979b. Winter nurogen torage m five rnacroaigae In 'arr3jU1ruClt Bay,R.1. J. Phycol. 15(Suppl ):15
1980. Arumal waste as a nitrogen source for Gra ,lanD IlkvahlOe and .....~
a/?ardh,ella batley, \0 culture. Aqu",-ulture 21 ~ 7·91
AUSTIN, A, and R. ADM1S
1974. Red algal resource studies to Cana(ban PacifIC walers. Fed . ,'>1mlSt
fj,h. Pro' ~1101St. Ra:reauon c.ons.=rv, nnu Rep. (1973), 256 p.
AUSTIN, A , R. ADAMS, '\ JONES, and h A!"DERS
1973. Development of a method for u"eymg red algal resources to Una
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1960. Obsenations on furcel/ana /asllfilOtl1 (t ) Lam. forma aegaf(IOplJ
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BARILOTTI, C. D., and W IL VERTHOR
1972. A resource management tud)· of Gel,d,um robwlum In K
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OJ' Tol.}o Press
Japan .
BARILOTII, D. O.
1974. Environmental factors that affe.'t the dl5tnbution and abundance 0
Geltdium robustum. In Scienufic Programme and Abstracts of Papers ,VlIlIntemationa! Seaweed SympoSIUm, Bangor, Wales. ug. I -24. 1974
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BELEAU, M. H., N. D . HElDELBAUOH. and D . VA DYKE.
1975. Open-ocean fanning of kelp for conversion to animal and humanfoods. Food Technol. 29(12):27-30, 45
BIRD, C. J.
1973. Aspects of the life history and ecology of Porph>ra lmeans(Bangiales, Rhodophyceae) in nature. Can. J . Bot. 51:2371-2379.
BIRD, C. J ., L. C.-M. CHEN, and 1 McLACHLAN
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BIRD, C. J ., T . EDELSTEIN. and J. McLACHLAN
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reproduction in Pomquet Harbour, Nova Scotia.
104:245-255 .
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BIRD, K. T.
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c) There is a continuing need for the support and trainingof talented phycological manpower (students) who willbe the seaweed farmers of the future.
d) An enchancement of scientific information (Le., exchanges) with Asiatic phycologists-aquacuJturists issimilarly important in the development of current andfuture seaweed maricuJture programs.

ACKNOWLEDGMENTS
The author would like to thank his wife, Myla Mathieson, for
her helpful editing of the manuscript. Donald Cheney andClayton Penniman of the Jackson Estuarine Laboratory andNortheastern University, respectively, are acknowledged for theircritical reviews and constructive comments on the paper. The typing skills and efforts of Debbi Jackson in the fmal completion of
the manuscript are gratefully acknowledged.

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19'4
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