Techniques in Finfish Larviculture in Taiwan

Aquaculture 200 Ž2001.
1–31
www.elsevier.comrlocateraqua-online
Techniques in finfish larviculture in Taiwan

I Chiu Liao a,) , Huei Meei Su b, Emily Y. Chang a
a
Taiwan Fisheries Research Institute, 199 Hou-Ih Road, Keelung 202, Taiwan
b
Tungkang Marine Laboratory, Taiwan Fisheries Research Institute, Tungkang, Pingtung 928, Taiwan
Abstract
Taiwan now has over 90 finfish species in which larviculture is possible. Billions of finfish fry
can be produced annually. The use of modern and advanced techniques in larviculture has won
Taiwan a leading position in the world. This achievement can be attributed to: Ž1. successful
broodstock management, including broodstock collection and cultivation, maturation and spawn-
ing, and egg collection and incubation, Ž2. complete larval rearing using outdoor and indoor
systems, and Ž3. establishment of techniques of live food preparation for larval feeding. Cannibal-
ism, difficulties in water quality control, and disease outbreak often cause mass mortality in the
rearing larvae. However, the obstacles that hinder the development of larvae at their later stages
are being overcome. Through a systematic understanding of the complex behavioral patterns of
finfish larvae, cannibalism can somewhat be controlled under larviculture conditions. Control
strategies often involve physical manipulations such as grading and feeding adjustments. For water
quality and disease control, an indoor larval rearing system has recently been established. It adopts
ozone or chlorine to disinfect fertilized eggs, the rearing water, and facilities. Additionally,
utilization of the green water technique in rearing larvae and of ozone-disinfected recirculating
water in weaning larvae has improved the hygiene of the systems. For instance, the average
survival rate in the fry of grouper used to be less than 0.1% until they attained a size of about 2
cm. With the improved system, grouper fry grow from 3 to 10 cm with almost no mortality or
viral infection, with a feed conversion rate of 0.7–0.9 and a daily weight increase rate of 4–27%.
Application of live food enrichment techniques with selected nutrients for short and long terms
has proven beneficial to larval growth. Studies determining the optimal environmental require-
ments of larvae have been essential towards successful larviculture. This paper mainly discusses
)
Corresponding author. Fax: q886-2-2462-4254.
E-mail address: icliao@mail.tfrin.gov.tw ŽI.C. Liao..
0044-8486r01r$ - see front matter q 2001 Published by Elsevier Science B.V.

PII: S 0 0 4 4 - 8 4 8 6 Ž 0 1 . 0 0 6 9 2 - 5
2 I.C. Liao et al.r Aquaculture 200 (2001) 1–31
developments and advanced techniques of finfish larviculture in Taiwan. q 2001 Published by

Elsevier Science B.V.
Keywords: Techniques; Finfish; Larviculture; Taiwan
1. Introduction
The practices of finfish culture in Taiwan can be traced back to the 15th and16th
centuries when emigrants from mainland China came to settle on the island ŽLiao,
1993.. Techniques of larviculture have gradually been developed from simply collecting
the stocking material in the wild, to holding them for further use, to growing them in a
pond, and to using modern, advanced facilities and treatments for complete larviculture
practices. In Taiwan, it was not until the early 1960s that modern techniques in
larviculture began to evolve. A major success was the establishment of techniques for
freshwater finfish propagation in grass carp Ž Ctenopharyngodon idellus . and silver carp
Ž Hypophthalmichthys molitrix . ŽLiu, 1963, 1964; Tang et al., 1963; Lin, 1965.. Follow-
ing that, propagation techniques for marine finfish were established for grey mullet,
Mugil cephalus ŽLiao et al., 1969.. With techniques developed for these species as the
benchmark, larviculture for over 90 finfish species is now possible in Taiwan ŽTable 1a
and b..
Many fisheries resources and wild fry have been seriously depleted due to overfishing
and environmental pollution. The availability of quality seedstock has been one of the
most critical factors in the commercial success of industrial production of finfish that
has stimulated continuous developments of finfish larviculture in Taiwan. In other parts
of the world, large-scale production of marine finfish fry began in the 1980s and has
continued onwards ŽSorgeloos et al., 1995., During this time, Japan pioneered the red
sea bream Ž Pagrus major . and Europe developed the most competitive hatchery
methods for the sea bass Ž Dicentrarchus labrax . and the gilthead sea bream Ž Sparus
aurata.. To assure a steady protein supply and good quality aquatic food for human
consumption, reliable and sound techniques in larviculture have become essential in
aquaculture now and for the future.
A complete finfish larviculture generally involves three phases of techniques, includ-
ing Ž1. broodstock management, Ž2. larval rearing, and Ž3. food preparation. Broodstock
management involves techniques to collect and culture high quality broodstock, to
induce maturation and spawning, to manipulate sex, to preserve sperm, and to collect
and incubate eggs. In larval rearing, techniques that have been developed concern the
larviculture system, nutrient requirements, cannibalism prevention, and pond manage-
ment. Nutritional content and size of food for larvae and fry determine the success of
commercial-scale larviculture. Food preparation for larviculture involves the use of
highly unsaturated fatty acids ŽHUFAs.-supplemented feeds, the application of suitable
live food for different larval stages, and the adoption of live food enrichment protocols.
Each time problems occur during the course of research and development of finfish fry
production in one target species, techniques applied to all these three phases must be
evaluated and modified. This paper describes techniques that have been successfully
I.C. Liao et al.r Aquaculture 200 (2001) 1–31 3
developed and are continuously being applied to finfish larviculture in Taiwan, specifi-
cally using freshwater finfish, milkfish Ž Chanos chanos ., groupers, cobia Ž Rachycentron
canadum., and eel Ž Anguilla japonica. as examples.
2. Broodstock management
The purpose of broodstock management is to supply good quality eggs and larvae. It
generally covers the collection and domestication of broodstock, maturation and spawn-
ing, and egg collection and incubation.
2.1. Collection of broodstock
A successful hatchery operation is based on healthy broodstock. Broodstock are

usually caught from the wild, bought from commercial grow-out ponds, or raised in the
hatchery for artificial propagation purposes. Different species require different ways of
collection and cultivation. In the past, broodstock of grey mullet were captured from the
wild during their spawning migration. Spawners of grey mullet were caught using an
outboard-powered raft, then brought into indoor tanks and immediately induced to
spawn by hormonal treatment ŽLiao et al., 1972; Liao, 1975.. Milkfish broodstock, on
the other hand, are rarely caught from the wild or are not available from commercial
grow-out ponds because at market size milkfish are generally too young for induced
maturation. Milkfish do not become sexually mature until many years after reaching the
market size. Therefore, young milkfish are usually raised in the hatchery’s grow-out
ponds, which may take at least 5 years ŽLiao and Chen, 1979..
For grouper fry production, collection of broodstock is the first bottleneck because
the mature fish are quite expensive and less available in captivity. Fish of maturing age
are difficult to obtain in large numbers. Groupers are protogynous hermaphrodites, being
female in their early life and turning into male at older ages, so that the males are
usually larger in size and rare in the population. Broodstock can be obtained from the
wild or from ponds, where young groupers are reared until sexual maturity. Pond-reared
broodstock have better survival rate and breeding performance than wild-caught brood-
stock.
2.2. Feeds
To assure good quality eggs, dietary lipid composition in broodstock feed is

important. The quality and quantity of n y 3 HUFAs contained in fish feeds may
influence development of the gonads and quality of the egg ŽNavas et al., 1998; Sargent
et al., 1999.. Recent studies have concentrated on the lipid content of feeds for
broodstock. Broodstock of seabass Ž D. labrax . that had insufficient feed or diet
produced poor quality eggs with a low fertilization rate ŽKatavic, 1986.. In Taiwan,
broodstock of carnivorous fish are generally fed with fresh trash fish Žmostly Scomber
australasicus.. Fresh trash fish has recently been found to improve gonadal develop-
Table 1
Ža. List of freshwater finfish larvicultured in Taiwan Žmodified from Liao, 2000.
Scientific name Common name Year of first success
Aristichthys nobilis Bighead carp 1963
Bidyanus bidyanus Silver perch 1994
Carassius auratus Gold crucian carp –
Channa maculata Snakehead 1970
Cirrhina molitorella Mud carp 1963
Clarias batrachus Thailand catfish –
C. fuscus Walking catfish 1970
Colossoma macropomum Pacu 1984
Ctenopharyngodon idellus Grass carp 1963
Culter erythropterus Redfin culter 1974
Cyprinus carpio Common carp –
Hypophthalmichthys molitrix Silver carp 1963
Lates calcarifer Giant perch 1983
Lateolabrax japonicus Japanese sea bass 1982
Leptobarbus hoeÕenii Sudan fish 1984
Megalobrama amblycephala Wu-Chang fish 1982
Micropterus salmoides Largemouth bass 1983
Misgurnus anguillicaudatus Pond loach 1977
Morone saxatilis= M. chrysops Hybrid striped bass 1995
Mylopharyngodon piceus Black carp 1966
Onchorynchus mykiss Rainbow trout 1964
Oreochromis aureus Blue tilapia 1975
O. mossambicus Mozambique tilapia 1946
O. mossambicus= O. niloticus Fu-So tilapia 1969
Oreochromis niloticus Nile tilapia 1967
O. niloticus= O. aureus Hybrid tilapia 1974
Oxyeleotris marmortus Marble goby 1973
Pangasius sutchi Thailand catfish 1976
Parasilurus asotus Chinese catfish 1971
Plecoglossus altiÕelis Sweet-fish 1979
Siniperca chuatsi Kuei Fa bass 1987
Varicorhinus barbatulus Taiwan shoveljaw carp 1986
Žb. List of marine finfish larvicultured in Taiwan Žmodified from Liao, 2000.
Acanthopagrus australis Australian sea bream 1979
A. berda Grey fin sea bream 1979
A. latus Yellowfin sea bream 1980
A. schlegeli Black sea bream 1979
A. siÕicolus Southern black sea bream 1980
Argyrosomus japonica Silver croaker 1994
Caranx ignobilis Goyan fish 1998
C. lugubris Giant jack 1999
C. melampygus Yellow fin jack 1999
C. papuensis Brassy trevally 1998
Chanos chanos Milkfish 1978
Choerodon schoenleinii Green wrasse 1998
Cromileptes altiÕelis Humpback grouper 1998
Eleutheronema tetradactylum Four finger thread fin 1996
Table 1 Ž continued .
Žb. List of marine finfish larvicultured in Taiwan Žmodified from Liao, 2000.
Scientific name Common name Year of first success
Epinephelus coioides Orange-spotted grouper 1986
E. fuscoguttatus Tiger grouper 1996
E. lanceolatus Giant grouper 1997
E. malabaricus Malabar grouper 1982
Glossogobius giuris Flathead goby 1990
Gnathanodon speciosus Golden fish or king fish 1998
Hapalogenys nitens Black grunt 1998
Larimicthys crocea Large yellow croaker 1997
Leiognathus equulus Common slipmouth 1998
Lethrinus haematopterus Common lenjan 1994
Lethrinus nebulosus Blue emperor 1983
Liza macrolepis Big scale liza 1997
L. subÕiridis Red eye liza 1997
Lutjanus argentimaculatus Red snapper 1984
L. erythropterus Pink snapper 1998
L. johnii Golden snapper 1997
L. monostigma One-spot snapper 1998
L. riÕulatus Yellowfin snapper 1998
L. russelli Russel’s snapper 1998
L. sebae Seba’s snapper 1998
L. stellatus White-spotted snapper 1998
Mugil cephalus Grey mullet 1969
Nibea diacanthus Spotted croaker 1991
Opleganthus punctatus Rock porgy 1994
Pagrus major Red sea bream 1979
Platax orbicularis Round batfish 1994
Plectorhinchus cinctus Yellow spotted grunt 1990
P. cinctus Žalbino. Three-banded sweetlip grunt 1997
P. diagrammus Stripe grunt 1996
Plectropomus leopardus Coral grouper 1989
Polynemus plebeius Striped threadfin 1995
Pomadasys kaakan Silver grunt 1990
Rachycentron canadum Cobia 1994
Scatophagus argus Spotted scat 1997
Sciaenops ocellatus Red drum 1991
Seriola dumerili Great yellow tail 1996
Siganus fuscescens Doctor fish 1996
S. guttatus Star siganus 1996
S. oramin Net-pattern spinfoot 1982
Sillago sihama Sand border 1992
Sparus sarba Silver sea bream 1984
Takifugu rubripes Tiger puffer 1994
Therapon jarbua Three-stripped tiger fish 1989
Trachinotus blochii Long dorsal fin pompano 1989
T. falcatus Permit fish 1989
T. oÕatus Short dorsal fin pompano 1997
–, cannot be determined.
ment, fertilization rate, and quality of fertilized eggs in seabass ŽNavas et al., 1998.. It
was speculated that the composition of the lipids in the feeds was the major factor for
the gonadal development in seabass. Formulated pellet feeds are generally used for
broodstock of omnivorous fish such as milkfish ŽChang et al., 1993.. Moist pellet feeds
can be used as carriers of extra nutrients such as vitamins, fish oil and some prophylac-
tics.
2.3. Sexual maturation and spawning
Spawning of broodstock could occur artificially through hormonal induction or

spontaneously through sophisticated practices of manipulation. Many fish, if properly
induced, will spawn under controlled conditions. The induction method that has been
popularly used in Taiwan for more than four decades is through the injection or
implantation of hormonal materials ŽTang et al., 1963; Liao, 1975; Liao et al., 1979;
Huang et al., 1986; Lin et al., 1986; Ho et al., 1997.. The fish, however, often incur
physiological stresses and injuries during induction processes, particularly during strip-
ping, and this made them more susceptible to diseases or even death. Fertilization and
hatching rates of eggs from induced spawns are generally low. The methods of
spontaneous spawning require the use of well-balanced and sound approaches to
cultivate the broodstock in captivity. Recently, hatchery operators and researchers have
tended to use the spontaneous spawning method because of its many advantages ŽTable
2..
During artificial propagation, handling of fish that are hermaphrodites is difficult
because fish have grown bigger and heavier when they complete sex reversal. Further-
more, it takes longer to culture the sex-reversed ones. Therefore, attempts were made to
manipulate sex in small young finfish. In grouper Ž Epinephelus malabaricus, formerly
E. salmonoides., females over 1 year old could be induced to sex reversal with oral
application of 17a-methyltestosterone at the dosage of 1.5 mgrkg body weight ŽBW.
over a 2-month duration ŽYeh et al., 1987. or by implantation of pelleted androgen ŽYeh
et al., 1988.. Sex-reversed females did not exhibit female characteristics and their genital
pore was closed. However, sex-reversed females produced a small quantity of sperm.
Convenience of artificial propagation will be enhanced if sex-reversal techniques are
further improved.
Table 2
List of advantages of spontaneous spawning in finfish larviculture Žmodified from Liao, 1993.
Advantages Remarks
Energy saving Ø More efficient than the conventional method.
Ø No need for procedures of induced maturation.
Ø Use of physiological, nutritional, ecological and endocrinological approaches.
Spawner saving Ø One spawner can be used several times due to reduced injuries.
Automation Ø Convenience in collecting, washing, and transferring of eggs.
Efficiency Ø Increased fertilization and hatching rates.
Quality control Ø Higher percentages of healthy fry Žin large-scale..
Artificial propagation is further assured with gamete cryopreservation. In Taiwan,

cryopreservation technology has been established in some cultured finfish and shellfish
ŽChao and Liao, 1999.. Materials that have been successfully cryopreserved include
sperm and embryos. This technology is especially useful in protogynous hermaphrodites
in which older and larger males are difficult to obtain. The utilization of cryopreserva-
tion at the farm level, however, is still limited.
2.4. Egg collection and incubation
Spawned eggs in broodstock ponds are generally collected with a trap net utilizing
the water current produced by a water paddlewheel ŽChang et al., 1993.. High densities
of diatoms and copepods, and suspended detritus, however, reduce the efficiency of egg
collection. Copepods can be separated from eggs by using a grading apparatus and used
as live food for finfish larvae. Filamentous macroalgae can be removed by stocking
some herbivorous species Že.g. rabbitfish, Siganus spp. and spotted scat, Scatophagus
argus . in the pond. In some ponds where small shrimp and goby are present, it is
important to clean, disinfect, and dry the pond before spawners are stocked. For milkfish
cultivation, carnivorous bottom fish such as the yellow spotted grunt Ž Plectorhinchus
cinctus . are recommended to be stocked in the spawning pond for the removal of tiny
shrimps and fish ŽChang et al., 1993..
Physical damage to eggs is another factor that affects egg collection. Recent studies
have tried to identify the sensitive stages of eggs in order to avoid unnecessary
mortality. Grouper eggs and newly hatched larvae, for instance, are very sensitive to
stress and handling ŽCaberoy and Quinitio, 1998.. Mortality is minimized by handling
only neurula-stage eggs Žafter the formation of the optic vesicles. or by stocking eggs
into the culture tanks 2 h before hatching so that handling of larvae is avoided ŽLim,
1993; Tamaru et al., 1995; Caberoy and Quinitio, 1998.. Newly hatched larvae are
sensitive to light and are generally kept under dark conditions.
There are generally two kinds of facilities that are used for egg incubation. One is the
0.5-tonne Fiberglass Reinforced Plastic ŽFRP. tank with a conical bottom and the other
is the rectangular net hapa Ž2 = 1 = 1 m.. In the tank system, both raceway and static
types are used. In the static-type tank, dead eggs that sink to the bottom are periodically
removed by draining the outlet while temporarily stopping the aeration. Usually, the
process of transferring eggs three to four times to different tanks of fresh seawater is
needed before hatching or stocking. For the raceway type, incubation is in a net of
columnar shape, suspended in a well-aerated tank. Stocking density of eggs should be
less than 1 kg per tank in the static system, but can be several times higher in the
raceway system. The best method used so far has been the type with the rectangular net
suspended in a concrete pond and with a continuous flow of fresh seawater.
The traditional method for collecting eggs has a trap net set at the outflow of the
spawning tank. Recently, egg collection and incubation have been combined and
automated. The automated larviculture production system, which is operated from egg
collection to hatching and transfer of larvae, consists of a spawning tank, a filter, an
incubation tank, hatching nets, and larviculture ponds ŽLiao, 1996; Liu et al., 2000.. In
comparison with the traditional method, this system produces higher rates of fertilization
and hatching. The use of filtration and recirculation of culture water makes the system
environment-friendly. Aside from being labor and cost saving, this system offers an
improvement to the finfish hatchery technology.
3. Larval rearing
The objective of larval rearing is to mass-produce healthy seed. Management of
rearing environment and the feeding regime are two most important aspects of this
activity. To improve larval rearing techniques, a good understanding of larval morphol-
ogy as well as larval behavior and reactions of larvae to feeds and environmental
conditions is fundamental.
3.1. Rearing systems
In Taiwan, larval rearing is operated by indoor and outdoor systems. Most of the
indoor systems for larval rearing use concrete tanks, which have depths of 1–2 m and a
volume of less than 100 m3. The outdoor system is usually bigger, with areas larger than
100 m2 and depths of 1.0–1.5 m, and has a lower operating cost. The indoor system is
only used for high-priced fish, such as grouper. A comparison of indoor and outdoor
systems is made in Table 3.
Fertile water, derived from the extracts of shrimp flakes, is often used in the
beginning of larval rearing to initialize algal growth. When larvae begin to prey on
copepods, fertile water is replaced gradually by clean water. Some fish like red snapper
Ž Lutjanus argentimaculatus. and cobia Ž R. canadum., in which the first feeding larvae
require special feeds, can only be cultured in the outdoor system. In an outdoor system,
a large pond of 3000 m2 Ž3000–5000 m tonnes. is good for easy operation. In these
ponds, food sources are sufficient, operating costs are low, and water quality does not
deteriorate easily even if the ponds have earthen bottoms; however, cannibalism and
predation by birds cause problems. In small outdoor ponds Ž300–600 tonnes., however,
water quality is unstable and variable and may cause larval mass mortality. Fry are
easily infected by disease from physical injury during harvest from a large pond. For
milkfish, the outdoor larval rearing system with an area of 200–300 m2 is equipped with
a black plastic shade to improve sunlight insulation and prevent excessive algal bloom.
Table 3
Comparison of indoor and outdoor larval rearing systems ŽLiao, 1996.
Characteristics Indoor Outdoor
Tankrpond water depth 1.0–2.0 m 1.0–1.5 m
Water volume -100 tonnes )100 tonnes
Larval survival rate High Unstable
Feed supply and water control Poor Easy
Larval growth Slow Fast
Fry quality Poor Good
Production cost High Low
Filamentous algal growth Impossible Possible
Since 1994, when mass mortality due to viral infection ŽNNV or nervous necrosis
virus. occurred in 2-week-old grouper fry ŽChi, 1997; Chi et al., 1997., an indoor larval
rearing system was developed. Larviculture has been improved in terms of hygiene,
nutrition and techniques. Disinfection of eggs is essential since it considerably reduces
the risk of bacterial and viral transmission from broodstock to larval rearing tanks
ŽSalvesen and Vadstein, 1995; Fukuda et al., 1996; Tanaka, 1998.. Eggs are washed and
disinfected by UV- or ozone-treated water. All seawater and equipment are disinfected
by chlorine. Larval rearing is performed in batch culture using the green water
technique, i.e. by addition of the microalga Isochrysis. This technique has several
advantages in microbial and nutritional aspects ŽReitan et al., 1993; Su et al., 1997b..
Algae can be used as food of rotifers and to control water quality. Bacterial development
during hatching and enrichment in Artemia can be controlled through addition of
disinfected cysts or disinfectant-appended enrichment diets Že.g. DC DHA Selco.. At the
weaning stage, larvae are reared in a recirculation system consisting of a T-shape water
inlet, a waste removal settler, a submerged biofilter, a disinfection unit based on
ozonization, a unit for oxygen addition, water quality control units, and a computerized
water quality monitoring system. Unlike the NNV-infected fry reared in the traditional
system, fry of red-spotted grouper Ž E. coioides . and giant grouper Ž E. lanceolatus.
cultured in this system are healthy and their body surface color is not blackened.
Juveniles can grow from 3 to 10 cm with almost no mortality or viral infection, with a
feed conversion rate of 0.7–0.9 and a daily weight increase of 4–27% Žunpublished
data..
3.2. Nutrient requirement
Larval rearing generally fails, as in eels, because of a lack of knowledge about larval
food requirements. In marine finfish larvae, there are two general models for the uptake
of nutrients from the environment ŽOtake et al., 1993; Otake, 1996.. The first is the
typical type that utilizes plankton, such as oyster trochophores, rotifers, Artemia, and
copepods as food sources. The requirement for n3-HUFAs is critical for finfish larvae in
this group. Most aquaculture finfish, such as milkfish, sea bream and grouper, belong to
this category. These finfish are thought to be a more evolutionarily advanced species.
The second type passively takes up dissolved organic materials ŽPfeiler, 1986.. Most
finfish in this category belong to the superorder of Elopomorpha, including Elopiforms,
Anguilliformes, and Notacanthiformes, in which a leptocephalus form is present during
the larval stage. These finfish are thought to be more primitive in evolution. Eel, one of
the species in this category, has very mysterious larval stages and its nutritional
requirements are barely known.
3.3. Cannibalism
In hatchery rearing conditions, cannibalism often occurs during the larval stage and it
affects fry production to a certain degree. Cannibalism during larval and early juvenile
stages, and its possible causes, has been documented in many fish species ŽTable 4.. It is
generally related to genetics and larval behavior ŽHecht and Pienaar, 1993.. Difference
Table 4
Cannibalism in finfish larviculture
Species Causative Cannibalistic conditions References
factors
Centropomus Size variation Ø Increasing predator:prey size ratios beyond. Dowd and
undecimalis 2:1 greatly increased cannibalism; cannibalism Clarke Ž1988.
ŽSnook. rates did not peak until juveniles were 16–18
mm; decreased rapidly with increasing size.
Density Ø Increasing density did not affect
cannibalism rates.
Channa striatus Size variation Ø Size differences increased the rate of Qin and
ŽSnakehead. cannibalism. FastŽ1996.
Dry food Ø Increased availability of formulated feed
reduced cannibalism.
Cyprinus carpio Density Ø The rate of cannibalism was positively Van Damme
ŽKoi carp. density-dependent in larvae and juveniles, et al. Ž1989.
which were fed only dry foods.
Esox lucius L. Size variation Ø Strong differences of size are considered Gres and
ŽNorthern pike. to be stimulants for cannibalism. Belaud Ž1995.
Live food Ø Continuous supply of live prey did not
prevent its early cannibalism.
Gadus morhua Size variation Ø Cannibalism occurred when the Folkvord Ž1991.,
ŽAtlantic cod. maximum:minimum weight ratio exceeded Folkvord and
3.5:1 and was the main mortality factor Ottera Ž1993.
when the ratio was above 8.5:1; higher losses
in the 0.2-g group than in the 0.8-g group.
Live food Ø While fed live zooplankton, ungraded
groups of 0.2- and 8-g cod could survive
93.5–97% in 4 weeks; cannibalism
among starved 0.2-g cod ceased after
addition of live zooplankton.
Starvation Ø Cannibalism was higher among starved
0.2-g cod than fed 0.2-g cod.
Perca fluÕiatilis Size variation Ø Intra-cohort cannibalism occurred among Brabrand
ŽPerch. body size 10.5"0.26 mm; consumption Ž1995., Melard
rate increased exponentially with size et al. Ž1996.
of predatory perch; cannibalism was
suppressed via size-sorting after 24 days
of rearing.
Live food Ø Cannibalism was reduced when
Daphnia pulex were added.
Density Ø Cannibalism was found most intense at
intermediate densities.
Sciaenops Size variation Ø Increasing predator: prey size ratios Dowd and
ocellatus beyond 2:1 greatly increased Clarke Ž1988.
ŽRed drum. cannibalism.
Density Ø Increasing fish density from 0.05 to 0.1
preyrl increased cannibalism from a mean
of 1.0 to 3.7 preyrpredatorrday.
Scomber scombrus Live food Ø Cannibalism declined with increasing Fortier and
ŽAtlantic mackerel. density of copepod nauplii in the Villeneuve
environment and with increasing number of Ž1996.
alternative copepod prey ingested.
Table 4 Ž continued .
Species Causative Cannibalistic conditions References
factors
Density Ø Cannibalism by Atlantic mackerel larvae
Ž3–14 mm long. does not appear to
contribute to their density-dependent
regulation.
Sparus Turbidity Ø Using Chlorella at 0.8–1.0=10 6 cellsr Li et al.
macrocephalus ml would effectively prevent sibling Ž1992.
ŽBlack sea bream. cannibalism.
Stizostedion Density Ø As fish densities increased, natural deaths Zakes Ž1997.
lucioperca increased but losses due to cannibalism
ŽEuropean pikeperch. decreased.
Theragra Size variation Ø Cannibalistic individuals could consume Sogard and
chalcogramma fish close to the maximum size physically Olla Ž1994.
Žalleye pollock. possible under gape limitation; cannibal
to prey size ratio was 1.7.
Ø Smaller pollock could be attacked to
cause mortality by larger pollock.
in size within a cohort due to genetic variation is a principal cause. On the other hand,
larval behavior is governed by environmental conditions, such as food availability, food
type, nutritional composition of the food, population density, light intensity, refuge
availability, and water clarity. Methods to minimize cannibalism include satiation
feeding, optimal feeding frequency, even distribution of food, live food supplement,
optimum particle size of the dry feed, determination of photic preferences, size grading
of larvae, removal of dominant cannibals, and determination of optimal stocking density.
In a recent study on the red drum Ž Sciaenops ocellatus ., size difference in tested
larvae was found to be the most important factor affecting cannibalism, while water
turbidity and larval density showed no obvious effects. Satiation feeding significantly
decreased, but did not stop, cannibalism. The peak stage of cannibalism for the larvae
was estimated to be at a total length of 1.8–1.9 cm, when the ratio of lower jaw length
to total length was the highest Žunpublished data.. Larval cannibalism was also found in
grouper. Yeh et al. Ž1996. investigated the effect of body length on cannibalism in the
orange-spotted grouper Ž E. coioides . and found that a total length of less than 6 cm to
be the peak stage of cannibalism in this fish. For the cannibalistic behavior to occur, the
total length of predators and prey forms a linear logarithmic relationship. In addition, a
formula for estimating the probability of cannibalism between predators and prey was
derived from their total length, in which size difference increased with the probability of
cannibalism ŽYeh et al., 1996..
3.4. Pond management
Two of the most important strategies in pond management are water quality control
and grading. In the former, addition of green algae Žnatural blooms or cultured
Nannochloropsis oculata. to rearing ponds and control of water exchange rate are the
main approaches. In the outdoor system, pond management is focused mainly on
controlling the stability of green water. Green water results from the growth of green
microalgae and functions in Ž1. stabilizing of water quality, Ž2. being utilized by live
food organisms, and Ž3. controlling the transparency to stabilize larvae. High trans-
parency is not favored for larval stability because it promotes growth of filamentous
macroalgae, which impede larval swimming. Diatom blooms, especially the develop-
ment of those with long spine, are harmful to larval survival. Monoculture of microalgae
is generally difficult to maintain in the outdoor rearing ponds; therefore, water that has
the dominant green algae is generally delivered from nearby grow-out ponds to the
rearing ponds ŽLiao, 1996.. The introduction should receive extra care because a change
of algal flora will lead to the mortality of larvae. In the indoor system, water quality is
controlled by: Ž1. addition of cultured algal water andror fermented water, which is
derived from trashfish and other fish by-products and extract from crushed shrimp
flakes; Ž2. removal of waste with a siphon; Ž3. partial or continuous water exchange; and
Ž4. appropriate adjustment of light intensity and temperature.
Grow-out farmers usually avoid disparity in fry growth, which may lead to cannibal-
ism and, thus, to a low survival rate. Therefore, fry are generally graded several times
before being shipped to customers. The grading apparatus mainly consists of stainless
wire with different mesh sizes. During the grading process, injury and subsequent
infection with pathogens may cause high mortality of fry.
4. Food preparation
Food preparation should ensure steady supplies of nutrition during larval rearing. It
includes techniques of live food culture, food enrichment, and artificial feed develop-
ment. Stability and reliability of live food culture are indispensable. Food of optimum
size, amount and quality should always be available to the larvae. Foods that are not
sufficiently nutritious for the larvae should be enriched with nutrients prior to feeding. In
Taiwan, selection, collection, and culture of live foods to be used in larviculture began
in 1982 ŽSu et al., 1997a.. As of 1997, there were 31 species Ž49 strains. of microalgae,
3 species Žnine strains. of rotifers, 1 cladoceran Ž Diaphanosoma aspinosum. and 1
copepod Ž Apocyclops royi . that were being held as starter cultures ŽSu et al., 1997a..
Foods that are optimal for larviculture include algae, oyster trochophores, rotifers,
copepods, nauplii and adult Artemia, cladocerans, red worms, and formulated feeds. A
combination of live foods yields better results in growing larvae. For instance, larvae of
the grouper Ž E. coioides . had better growth and survival rates since F1 Žfirst feeding
day. when subsequently provided with oyster trochophores ŽF1–F3., SS-rotifers ŽF1–F6.,
and S-rotifers ŽF7–F15. ŽSu et al., 1997b.. For marine finfish larviculture, food often
consists of microalgae, oyster trochophores, rotifers, copepods, and Artemia. Both
oyster trochophores and copepods are very nutritious larval foods, containing more than
10 mgrg dry weight ŽDW. of eicosapentaenoic acid ŽEPA. and docosahexaenoic acid
ŽDHA.. Furthermore, copepods have the highest DHA to EPA ratio. On the contrary,
rotifers and Artemia are poor in HUFA and need to be enriched before feeding.
In freshwater larviculture, green water and live food are cultured in the rearing ponds
prior to the release of fish larvae. Crushed steamed egg yolk, smashed red worm, and
copepods are often used as food for freshwater larval rearing. The cladocerans are often
used in the nursery stage of the Asian sea bass Ž Lates calcarifer .. Green microalgae that
are maintained with pig manure and ground soybean powder are commonly used as feed
for the cladocerans. Red worms are often used as food for eel elvers and larvae of
catfish Ž Clarias spp.. and largemouth bass Ž Micropterus salmoides.. Red worms are
usually collected from the bottom of small streams, or irrigation canals and ditches
where organic matter accumulates.
4.1. Microalgae
Finfish larvae take up microalgae passively with water or indirectly through live
food. The algae Isochrysis and Nannochloropsis are used in the first feeding due to
their high DHA and EPA content, respectively. Nannochloropsis, however, is not
suitable for the trocophore-feeding stage of the finfish larvae due to its indigestibility. N.
oculata, formerly known as marine Chlorella, is small in size Ž2–5 mm., but contains
30% EPA. It is a nutritious food for rotifers, but not suitable for bivalves, Artemia, and
copepods. The local strain of I. galbana contains about 12% or 10 mgrg DW DHA,
which is good for rotifers and Artemia ŽSu et al., 1997a.. In comparison with yeast,
bacteria, and formulated feed, microalgae are the best food for rotifers ŽHirayama et al.,
1979; Hirayama and Funamoto, 1983; Teshima et al., 1981; Su et al., 1998..
Microalgae are maintained in batch culture Ž20 8C, 20–60 mErcm2rs light, 12Lr12D
light period.. Seawater for general culture of microalgae is filtered through sand filter
and disinfected by hypochlorite, which is later neutralized by thiosulphate. Seawater for
stock culture is filtered sequentially through 1-, 0.8-, and 0.4-mm filters and autoclaved.
As to the culture medium, Walne’s Ž1974. formula is used for brown microalgae
Ž Isochrysis, Chaetoceros. in the indoor culture system Ž1, 10, 100 l., and agricultural
fertilizers are used for the outdoor culture system Ž500 l.. For green microalgae
Ž Nannochloropsis and Tetraselmis., green medium is used in 1- to 500-l culture ŽLiao et
al., 1991..
The nutritional content of microalgae varies with culture conditions, especially with
the growth stage ŽSu et al., 1988.. Generally, the optimum biomass and nutritional
quality can be reached during the late exponential growth stage. Thus, production
systems are designed to have several sets of modules, depending on daily demand. In
one module, there are different types of cultures, such as starter cultures, 1- to 3-day-old
cultures, and cultures ready for harvest. A successful culture of microalgae is obtained
by good quality inoculum, appropriate treatment of culture water, and regular renewal of
the culture.
4.2. Oyster trochophores
Oyster trochophores are widely used as live food in fry production of marine finfish,
especially in larvae that have a small mouth at hatching, such as the grouper. The HUFA
content of oyster trochophores is higher than that of the rotifers Žunpublished data..
Presently, oyster trochophores are often used for culturing grouper larvae. The survival
rate of grouper larvae is improved when oyster trochophores are added to the feed for 3
to 6 days after the first feeding ŽLin et al., 1986; Su et al., 1997b.. However, thorough
cleaning of the oysters is needed to prevent contamination of nematodes and parasites.
The quantity and frequency of feeding must be controlled to avoid deterioration of water
quality in the larval rearing tank.
4.3. Rotifers
Rotifers became one of the major larval feeds when Ito Ž1960. successfully applied
them to feed finfish larvae. Fushimi Ž1983. estimated the uptake of rotifers per fish larva
Žred sea bream. to be about 1.7–2.3 = 10 4 during its rotifer-feeding stage. In Taiwan,
rotifers are mostly mass-produced outdoors due to the lower operating cost. Outdoor
ponds are initially fertilized with chopped trash fish or scraps, such as entrails, gills, and
shrimp heads. Rotifers are then inoculated in the ponds. Recently, fermented water from
trash fish, fish soluble and photosynthetic bacteria have been appended to rotifer culture.
Rotifers are generally harvested in the early morning when they migrate to the upper
water column.
In Taiwan, three types of rotifers, i.e. L-, S-, and SS-type, or Brachionus plicatilis,
B. rotundiformis, and Brachionus sp., respectively, are found, with size range of
100–305 mm ŽSu et al., 1997a.. Most rotifers found in aquaculture ponds belong to the
S-type, which occurs year round, while L-type occurs only in the winter and SS-type in
the summer ŽSu et al., 1994a.. Temperature, salinity, and feed concentration all affect
the growth rate of the L- and S-types ŽLiao et al., 1991.. The most suitable temperatures
are between 28 and 32 8C. Above 28 8C, the effect of feed concentration on growth of
rotifer culture is more significant than that of salinity and size. Their growth rate
increases with decreasing salinity from 30 to 10 ppt. It is important that enough food is
provided during the warm seasons and that enough heat is provided during the cold
seasons. To feed rotifers, microalgae, bakery yeast, yeast powder, prepared feeds, and
chicken droppings are directly or indirectly used, but the best growth is obtained with
microalgae Že.g. Tetraselmis chui .. If heaters are not used during cold seasons, it is
preferable to culture large-type rotifers and lower the salinity.
As to the nutritional aspect, the HUFA content of rotifers varies a great deal and is
related to its feed. The EPA content in rotifers is highest Žabout 15–20%. when fed with
N. oculata, lower Žabout 5–10%. with Isochrysis aff. galbana or T. chui, and lowest
Žnearly 0%. with Chlorella sp. or yeast ŽSu et al., 1994b.. At present, most hatcheries
consider the algal species N. oculata as the best nutritive feed for rotifers. However, the
culture of N. oculata is very unstable and easily contaminated with protozoans.
Techniques that can depress or diminish protozoan contamination should be developed.
Recently, more stable and higher quality grouper fry are produced in the laboratory
using the Taiwan strain I. galbana tml as larval feed. I. galbana tml is the best among
the microalgae used in larviculture in terms of its DHA content, which is 10 mgrg DW
ŽSu et al., 1997a.. This new development has not been extended yet to the aquafarmers
because of the difficulty in the outdoor mass culture of the microalgae.
During the course of rotifer cultivation, there exist some unidentified causes of mass
mortality. Factors that may be involved include: Ž1. internal factors such as species-
specific requirements, and Ž2. environmental factors such as water quality, bacterial and
protozoan contamination. Some bacteria such as Pseudomonas acinetobacter facilitate
rotifer growth, but some pigment-producing bacteria such as FlaÕobacterium inhibit
their growth. Pseudomonas, which can produce vitamin B12 , is beneficial to rotifer
propagation ŽYu et al., 1988., but Vibrio alginolyticus can cause mortality of rotifers
ŽYu et al., 1990.. The influence of protozoa on rotifers is not well understood, but these
organisms probably compete for space, food, and oxygen ŽHino et al., 1981; Cheng,
1997.. Outdoor-produced rotifers may be contaminated with pathogenic bacteria, which
are harmful to finfish larvae. It is useful to apply antibiotics to control bacterial
contamination; however, drug-resistant strains and drug residues cause serious problems.
Recently, probiotic bacteria that have been applied by local farmers to modify the
aquaculture environment, might prove beneficial to rotifer cultivation. Probiotic bacteria
are speculated to improve water quality and inhibit the growth of harmful bacteria.
Furthermore, probiotic bacteria carried by rotifers may be ingested by the finfish larvae
and increase larval survival and disease resistance.
4.4. Copepods
During the late stage of larval rearing, copepods are important sources of live foods.
Copepod blooms usually occur in brackishwater ponds, but timing of the blooms for
larval feeding is difficult. Copepods are generally collected from fish and prawn
grow-out ponds. The collection and sale of copepods has become a profitable aquacul-
ture sub-business in Taiwan. Some farmers use big earthen outdoor ponds Ž1000–4000
m2 . to produce copepods. Copepods are fed with trash fish, fish powder and pellet feed
that are sprayed or soaked in the pond water.
The AcopepodsB used by private grouper hatcheries in indoor systems are actually the
copepods Schmackeria dubia Žcurrently known as Pseudodiaptomus annandalei . to-
gether with A. royi and the cladoceran Dia. aspinosum. Their size ranges are 250–1280,
110–1100, and 400–1150 mm, respectively. A. royi cultured in the laboratory fed with
microalgae develops from nauplius I to copepodite I in 5 to 6 days with a survival rate
of 89.6%, making it a highly potential species for mass production. In contrast, the
European copepod species, Tisbe sp., used for larviculture takes 12–25 days to reach the
copepodite I stage with a survival rate of less than 50%.
4.5. Artemia
Artemia nauplii are used as live feeds during the transition from rotifer to copepod
feeding. More uniform larval growth is achieved if Artemia nauplii are continuously
supplied for at least 2 to 3 days. However, contents of HUFAs in Artemia are
insufficient. Therefore, Artemia is often enriched with emulsified oil, which has a high
content of HUFAs and a suitable ratio of DHArEPArARA Žarachidonic acid..
The supply of adult Artemia is especially important to the finfish larvae during
metamorphosis. It reduces larval cannibalism and increases larval survival rate. Some
aquafarmers are specialized in the production of sub-adult Artemia. Ground rice bran
and fish powder feeds are generally used as food for adult Artemia. Adult Artemia
purchased from these farmers are transported in bags at water temperature of 14–16 8C
and fed as live food either directly or after enrichment with HUFA.
4.6. Others
Other foods used in seed production include shrimp flake, microparticulate feeds, eel
powder feeds, and steamed egg yolk and egg custard. Shrimp flake and microparticulate
feeds are also used to fertilize the rearing water. Sergestidae shrimp Ž Acetes inter-
medius . is an important dietary feed for fry after metamorphosis. The shrimps are
ground to different sizes according to sizes of fry, and then stored in a freezer. Frozen
ground shrimp can be fed directly to the fry.
Commercial feeds, on the other hand, are also available for fry production of marine
finfish. Formulated feeds larger than 1.5 mm are commercially available in Taiwan.
Artificial feeds with particle size larger than 2.5 mm are very popular. Weaning feeds
Ž- 1 mm. for grouper larvae are imported from Japan and Belgium and fed simultane-
ously with Artemia or copepods.
5. Case discussion
The following uses cases of several typical aquaculture species to demonstrate

techniques of larviculture currently practiced in Taiwan.
5.1. Freshwater finfish
Taiwan has a limited number of endemic freshwater species suitable for aquaculture.
Most cultured freshwater finfish have been introduced from foreign countries. Larvicul-
ture techniques for most freshwater finfish that have been established are based on those
that were developed for Chinese carps. Propagation techniques have been well devel-
oped for five Chinese carp species. These include grass carp Ž C. idellus ., silver carp Ž H.
molitrix ., mud carp Ž Cirrhina molitorella., black carp Ž Mylopharyngodon piceus ., and
bighead carp Ž Aristichthys nobilis .. Hatchery of Chinese carps in Taiwan is operated
from March to August. The first and most important step in the artificial propagation of
Chinese carps is the culture of broodstock, including selection, feeding, and transporta-
tion. Early on, fertilized eggs were mostly produced by hormone-induced strip spawn-
ing. An automated system that integrates spontaneous spawning, egg collection, hatching
and transfer of larvae has recently been developed and is now being routinely used
ŽLiao, 1996; Liu et al., 2000.. When larvae grow to 1 month old, they are ready for
stocking in the grow-out ponds.
Silver perch Ž Bidyanus bidyanus . was introduced from Australia in 1992. Using the
newly developed automated larviculture production system that is the same as that used
for Chinese carps, spontaneous spawning and mass production of silver perch larvae
were successfully achieved in 1994 ŽLiao, 1996; Liu et al., 2000.. The fish are induced
to spawn with a single injection of human chorionic gonadotropin ŽHCG. at 200 IUrkg
BW. Spawning seasons are normally from April to July. Fertilization rates of eggs reach
above 80% in May–July, but decrease to about 50% in August to September, and to 0%
thereafter. Larvae open their mouths and start to feed 48 and 96 h after hatching,
respectively, taking foods sequentially from rotifers, to cladocerans, to copepods, to
tubifix worms, and to aquatic insects.
Hybrid striped bass was introduced from the United States in 1992. It is a hybrid
between striped bass Ž Morone saxatilis . and white bass Ž M. chrysops .. In 1995,
spontaneous spawning in hybrid striped bass ŽF1 . was achieved. The F1 progeny were
cultured using commercial floating pellets. Two-year-old fish were already sexually
mature. Three-year-old females weighing 0.8–1.2 kg and males weighing 0.5–0.7 kg
were induced to spawn by a single hormone injection. HCG at a dosage of 300–400
IUrkg BW was the most effective treatment ŽLiu et al., 1998.. Induced fish were either
stripped or left to spawn in a tank. Hatched larvae were reared in 2 m3 circular ŽFRP.
tanks and fed sequentially with rotifers, Artemia, Daphnia, and fish starter crumble
feed. Larvae developed to the juvenile stage in 21–28 days after hatching, with total
length ranging from 12.1 to 18.2 mm. After 77 days of culture, juveniles reached a mean
total length of 7.5 cm with an average survival rate of 2.65% ŽLiu et al., 1998.. The F2
progeny exhibited low hatching and survival rates, slow growth, and high levels of
deformities and growth variability, suggesting that F2 progeny are not suitable for
aquaculture.
5.2. Milkfish (Cha. chanos)
Milkfish culture in Taiwan has a history of over 300 years. Its industry has now
evolved into a highly organized system with many specialized subsystems, including the
broodstock owner, the hatchery operator, the live and formulated feed supplier, the fry
broker and the harvester. The first success in artificial propagation of milkfish was
recorded in 1978 ŽLiao et al., 1979., and in spontaneous spawning in 1983 ŽLin, 1984..
These breakthroughs have increased and assured the supply of milkfish fry in Taiwan.
Historically, the demand for milkfish fry was more than 160 million per year, which
were either collected from the wild or imported from Southeast Asian countries. After
the breakthroughs, Taiwan has not only supported its domestic need of milkfish fry but
also exported them to other countries.
Like other marine finfish in Taiwan, the milkfish hatchery industry is usually
governed by two sub-systems. One is specialized in broodstock maintenance; the other
one is specialized in larval rearing. In the former, age and minimum body weight for
maturation, broodstock feed, and broodstock management require attention. It has been
reported that 5-year-old male and 6-year-old female milkfish become mature in captivity
ŽLiao and Chen, 1984.. However, satisfactory spontaneous spawning only occurred at
ages above 10 years ŽLin, 1984.. Wild milkfish have mature gonads at 2.5 kg, but
cultured ones do not have mature gonads when body weight is below 3 kg. For
commercial-scale breeding, spawners should be over 4 kg. Although spawning has been
achieved in captivity, the nutritional requirement of the broodstock is not well known
yet. At Taiwan Fisheries Research Institute ŽTFRI., a specially formulated feed consist-
ing of 70% formulated eel feed, 14.75% wheat germ meal, 14.75% soybean and 0.5%
vitamin E was prepared for the spawners during the spawning season ŽChang et al.,
1993..
For broodstock cultivation, milkfish that have not reached maturity are cultured in
low density with other main culture species, such as mullet, grouper and tilapia. Since
spawners with well-developed gonads are susceptible to handling stress, broodstock are
usually delivered to the spawning pond 2 months before the onset of spawning. To
stimulate feeding activities, stocking density can be increased in the winter. Gonadal
development during the spawning season is better in fish that were well fed in winter.
Handling of spawners is done very carefully because jumping often occurs and causes
injury. A 1-m-high net fence or fish nets are usually fixed along the dike to prevent
spawners from jumping out of the pond, which would happen with earthquakes, strong
light exposure, and high stocking density.
For milkfish larval rearing, outdoor ponds covered with black plastic shade are
generally used. The black covering, installed during the first 10 days after hatching,
prevents strong sunlight that can cause algal blooms and ultraviolet exposure, which are
harmful to the larvae. Most aquafarmers prefer the outdoor system because costs for
construction and fry production are low, management is simple, number of produced fry
is large, and larval growth is fast and healthy. The indoor system is much less favored
by hatchery operators because quality of fry produced indoors is generally unacceptable
for grow-out farmers. The optimum size for the outdoor pond is about 200–300 m2 and
1–1.2 m in depth. The pond bottom is covered with 30 cm of sand, which facilitates
water cleaning and harvesting. To ensure enough oxygen, the pond is aerated through
air–stones set 2.5 m apart from each other. Ponds are provided with individual inlets for
seawater, green water, and freshwater supply. Normally, 0.2–0.6 million larvae can be
produced in a 200–300 m2 pond.
A suitable stocking density of milkfish larvae at first feeding is about 2–3 larvaerl.
Larvae have strong phototaxis behavior. At 3–5 days old, the larvae tend to aggregate
and swirl at the corner of the pond, a dangerous behavior observed especially with
higher light intensity, higher water transparency and higher stocking density. Water
transparency can be controlled by adding green algae. Diatom and algal blooms,
however, cause bubble disease. To prevent this disease, it is necessary to control light
intensity, algal species, and algal density.
In the laboratory, milkfish larvae after hatching are fed sequentially with green algae
and cultivated plankton Žfrom Day 2., oyster trochophores ŽDays 3–8., rotifers Žfrom
Day 5., and white fishmeal and micropellet feeds Žfrom Day 15.. The supply of
cultivated plankton and oyster trochophores is important for increasing the survival rate
of milkfish larvae. Larvae can feed on smaller rotifers from the first feeding and addition
of oyster trochophores can improve larval survival rate ŽChang et al., 1993.. For mass
fry production, cultivated plankton and rotifers are used. Plankton is cultivated by
adding trash fish and powder feed into the rearing pond or into a separate plankton
cultivation pond.
Although techniques in milkfish larviculture have been well developed in Taiwan,
some problems have yet to be solved. Firstly, spawning Žfrom April. does not match
timing with high fry price ŽFebruary–May.. Therefore, manipulation of milkfish spawn-
ing in February or March is a challenge. Secondly, the cost of live larval food often
overwhelms the fry price. Thus, instead of oyster trochophores or rotifers, hatcheries
tend to cultivate cheaper live food Žcultured plankton., which has caused a gradual
decline in larval quality. Lastly, outdoor larval rearing facilities should be further
improved to compensate for unstable weather conditions. Heavy rainfall not only
induces collapse of algal blooms, but also causes poor water quality, which is harmful to
the larvae. In recent years, acid rain has become a problem.
5.3. Groupers
Groupers are among the most highly valued marine finfishes in Taiwan. Fry are
generally obtained from local hatcheries and wild fry are imported from Thailand, the
Philippines and Sri Lanka. The quality of imported wild fry is often poor, making
hatchery-reared fry more popular and expensive. Although more than 52 species of
groupers occur along the coasts of Taiwan ŽLiao, 1993., the most often propagated
species are the malabar grouper Ž E. malabaricus., orange-spotted grouper Ž E. coioides .,
tiger grouper Ž E. fuscoguttatus., and giant grouper Ž E. lanceolatus.. Among these, the
complete life cycle in captivity has been achieved in the orange-spotted grouper. Coral
grouper Ž Plectropomus leopardus. and humpback grouper Ž Cromileptes altiÕelis . are
propagated in small quantity.
Grouper spawners are cultured and maintained in earthen ponds or net cages, and
mature ones are selected for spawning. Cannulation of the spawner is needed to
determine the developmental stage of the egg. Previously, mature grouper spawners
were generally induced to spawn through injection and implantation of hormonal
materials ŽHuang et al., 1986; Lin et al., 1988.. Not until recently has spontaneous
spawning become successful in the grouper ŽLiao, 1993.. Groupers are protogynous
hermaphrodites, so male broodstock are difficult to obtain. Using sex reversal ŽYeh et
al., 1987, 1991. and cryopreservation techniques ŽChao et al., 1991., this problem can be
partially solved.
Collection of fertilized eggs is done with a trap net set at the outflow of the spawning
tank or downstream of a paddlewheel. Eggs are incubated in a suspended hatching net
with upflowing water. For transportation, eggs at the embryonic formation stage are
loaded at a density of 300 grbag in plastic bags filled with oxygenated seawater Ž10 l..
Salinity can influence the hatching rate of eggs. In malabar grouper, the hatching rate
varies from 14% to 76% in salinities ranging from 14 to 36 ppt. The hatching rate is
highest at 30 ppt.
Since 1985, larval rearing of the grouper has been studied in Taiwan ŽLin et al.,
1986.. Hatcheries for prawn post-larvae production were adapted and modified for
grouper fry production. These converted hatcheries were indoor systems with tank
volumes of 30–50 tonnes. Using indoor systems like this, fry survival rate could reach
as high as 30%. Mortality would occasionally occur in grouper fry of 0–7 and 18–19
days after hatching, and during metamorphosis. Mortality during the first stage Ži.e. 0–7
days after hatching. can be due to egg quality, food availability, and environmental
factors including water quality, aeration and light intensity; during the second stage Ži.e.
18–19 days after hatching., insufficient nutrients and viral infection cause mortality.
During metamorphosis, mortality is caused by deteriorating water quality, viral infec-
tion, and cannibalism. Generally, the survival rate of grouper larvae is very low
compared to other species ŽHussain and Higuchi, 1980; Fukuhara, 1989; Duray et al.,
1996, 1997; Kayano and Ho, 1997..
Larval mortality in humpback grouper Ž Cro. altiÕelis ., described in more detail from
a larval rearing experience in Indonesia, was characterized by six stages ŽTridjoko et al.,
1999.. The first stage is during incubation, when egg quality is not good and embryonic
development does not normally progress. The second, which has the most mortality,
occurs during the so-called floating death stage Ži.e. dead larvae float at the water
surface.. Egg quality, collision, and stress sensitivity are some influential factors. The
third is during first feeding, wherein some larvae are unable to switch from endogenous
to exogenous feeding. The fourth is in the 2-week-old larvae that are easily disturbed by
activities such as water exchange, addition of green water, and tank cleaning. The fifth is
in the 3-week-old larvae, in which gradual mortality is attributed to nutritional defi-
ciency. The last is in 4-week-old larvae when the larvae metamorphose, during which
cannibalism and nutritional deficiency cause larval mortality.
In Taiwan, mass mortality of grouper larvae is often caused by viral infection. Viral
nervous necrosis ŽVNN. disease occurs during hatching and larval rearing stages, while
iridovirus-like infection occurs during larval rearing and grow-out stages ŽChao and
Pang, 1997; Chi et al., 1997; Chi, 1997.. Larvae infected with the VNN virus, exhibit
unusual swimming behavior, such as whirling near the water surface, swimming
abruptly in a corkscrew manner, and sinking and floating periodically. In Taiwan, VNN
disease repeatedly occurs from April to September with peak mortality during the
summer Žfrom June to August., when the average temperature is around 30–32 8C. In
winter, the VNN virus can still infect the fish, but it does not cause cell necrosis or
mortality of the fish ŽChi et al., 1997.. In iridovirus-like infection, fry die gradually over
1 or 2 months from the onset of clinical symptoms. Fry and juveniles have a mortality
rate above 60%. Affected fish display blackening of the body surface, have breathing
difficulties and reduced feed uptake; however, no lesions are observed. Larval spleen
and head kidney swell 2 to 10 times compared to the normal size. In addition, gill
filaments are congested with subacute to acute petechia. Viral infections can be
controlled using ozone or chlorine to disinfect fertilized eggs, rearing water, and
facilities, and feeding the larvae with virus-free live food or artificial diets.
Comparatively, the survival rates in outdoor systems, adopted by hatchery operators,
are quite inconsistent. But occasionally, 500,000 fry in a crop are produced from a
3000-m2 outdoor pond. During autumn, the outdoor pond system benefits from the
temperate weather conditions. In general, larvae of better quality and price are produced
in indoor systems.
Giant grouper Ž E. lanceolatus. mass fry production was established in Taiwan in
1996 ŽHo et al., 1997.. Broodstock of the giant grouper mature and produce eggs in
concrete ponds. The spawners have initial body weights from 20 to 40 kg and can grow
to 40 to 110 kg when reared for 3 years in outdoor ponds Ž60 = 40 = 3 m.. Spawning
seasons last from May to October each year, with peak spawning from August to
September. Dosages of 390–400 IU HCG and 41–42 mg luteinizing hormone-releasing
hormone analog ŽLHRHa.rkg BW are used for injection in early May and June. A
dosage of 350 IU HCG and 38 mg LHRHarkg BW is used in the mid-spawning season.
Ovulation occurs monthly about 48 h after injection. In August of 1996 and 1997, the
total number of eggs produced was 28.8 and 195 million, respectively. Egg production
lasts for 1–3 days. Fertilized eggs are pelagic, transparent, individually separate, and
have a single centrally located oil droplet Ž0.18–0.19 mm.. Egg diameter ranges from
0.80 to 0.89 mm. Hatching occurs approximately 19.5 h after fertilization at water
temperature of 28–29 8C. The hatching rate is highest Ž90%. at 30 ppt. Eggs fail to
hatch at salinities lower than 20 ppt.
Newly hatched larvae have a total length of 1.65–1.91 mm and an oval yolk sac. The
oil droplet is located at the rear end of the yolk sac. The yolk-sac stage lasts for 2 days.
At the end of the yolk-sac stage, the average total length of the larvae is 2.91 mm. Since
hatching, the average total length of fry reared indoors have reached 4.0–5.5 mm by 15
days, 18.3–22.0 mm by 37 days Ži.e. adult-figured fry which can be harvested., and
20.7–32.3 mm by 46 days Ži.e. artificial diets can be used. Žunpublished data.. The
survival rate is higher than 90% when juveniles Ž) 2.6 cm. are reared in a recirculating
system and fed formulated feed until they reach 10 cm in total length.
There are some difficulties in the larviculture of giant grouper. Firstly, spontaneous
spawning is still not established and egg quality is unstable. Secondly, it is difficult to
obtain suitable live foods for early larvae. For first feeding, it is difficult to use rotifers
Ž150–250 mm., but is easy to use oyster trochophore Ž60 mm. and copepod nauplii Ž100
mm.. Oyster fertility, however, is highly variable, and techniques to separate and obtain
copepod nauplii are not yet developed. Thirdly, high water temperature that causes rapid
propagation of protozoans influences the stability of phytoplankton and thus water
quality, causing extremely high mortality in the early larvae Žbefore and during first
feeding.. Lastly, survival rate of 2-cm fry to 6-cm juvenile is low Ž- 30%..
5.4. Cobia (R. canadum)
Cobia, also known as sergeant fish, is a middle-sized tropical marine fish. It is

especially suited for sea cage aquaculture. Among the economically important species,
cobia is the fastest grower and most disease-resistant species. It has a long reproductive
period. Larviculture is not particularly difficult. High culture density is feasible. Fry can
be easily trained to take floating or sinking dry pellet feeds.
Unlike milkfish or grouper, 1-year-old cobia are already sexually mature. Cobia of
1.5 years spawns spontaneously. In southern Taiwan, the spawning season of cobia
peaks between February and May, and then periodically until October. Suitable water
temperatures for spawning are 24–29 8C, and for peak spawning are 24–26 8C. In
spring, spontaneous spawning behavior takes place at around 5 PM on a sunny day.
During summer, when daytime is longer, spawning is somewhat delayed until around
6–7 PM. Fertilized eggs float and are transparent, circular, and cream-colored. Eggs are
relatively large, with a diameter around 1.35–1.40 mm. Fertilized eggs hatch within 30 h
at water temperatures of 24–26 8C.
Newly hatched larvae have an average total length of 3.5 mm. The oil droplet is
located at the posterior end of the yolk sac. Black pigments are distributed all over the
body. Without air pumping, larvae stay in the water column with their ventral side up.
Larval growth is fast: 12 h after hatching they measure 4.0 mm in length; 12 h later they
have reached 4.73 mm. Cobia larvae generally have stronger vitality and tolerances than
other marine finfish larvae. Larvae of 3 days old can reach a total length of 5.1 mm.
First feeding starts on the third day, during which only rotifers and copepod nauplii are
fed. On the sixth day, larvae that have grown to 6.8 mm in total length are often fed with
adult copepods. Metamorphosis starts at around days 10–11. Larval color changes from
dark reddish brown to black, with the dorsal color changing to dark green. Fins become
colorful and stripes appear on the back. The period of metamorphosis is short, lasting
only about 1 day. After this, the larvae tend to stay at the bottom of the tank.
Size variation among larvae becomes significant 18 to 25 days after hatching. It is
good to collect and grade larvae at this stage to prevent cannibalistic behavior. Grading
is done every 3–4 days until larvae are 35–45 days old. Larvae less than 1 month old
are fed primarily with copepods. Larvae start to take Sergestidae shrimp or artificial
feeds after 1 month. Adult Artemia, if co-fed with copepods and pellet feeds, increase
larval survival rate, but increase the cost. Fry that have grown to 9 cm Žabout 45 days
old. are ready to be stocked into cage culture.
Some problems still exist in the larviculture of cobia. Size variation that leads to
larval cannibalism should be carefully studied. In addition to the use of strategies like
grading or increasing feeding frequency, optimization of larval feeds in size and kind is
one of the priorities for this species. On the other hand, the minimum space required for
larval rearing is not well known. In a space that is limited or small, cobia larvae
generally have low survival rate.
5.5. Eels
Eels are one of the economically important species in Taiwan. Since their artificial
propagation is not successful yet, glass eels for aquaculture are entirely being caught
from the wild. The unstable supply of glass eels has impeded the development of eel
aquaculture and related businesses. When Japanese eel larvae of 6 days old were first
successfully reared in the laboratory in 1974 ŽYamamoto and Yamauchi, 1974., the hope
for artificial propagation of eels was revived in the whole world. However, the duration
of larval survival has been poorly improved in the past decades, which may be due to
insufficient supply of good quality fertilized eggs, and lack of documentation on natural
foods and the biology of the larvae. There exist several stages in eel development from
larva to adult, i.e. from preleptocephalus, leptocephalus, glass eel, elver, yellow eel, to
silver eel. In the past two decades, eel larviculture has been focused on the first
developmental stage Ži.e. preleptocephalus.. Recently, the culture of larvae has made
significant progress, as the leptocephalus stage has been reached ŽTanaka, 1999a..
Eel broodstock can hardly be caught from the wild. Man-made obstacles like dams,
environmental pollution, and overcollection of glass eels have resulted in the scarcity of
wild broodstock. Therefore, pond-reared eels are utilized for artificial propagation
purposes ŽChang and Liao, 1977.. Due to poor knowledge on gonadal maturation and
gametogenesis, a lot of attention has been dedicated to artificial induction through
hormone treatments. In Taiwan, Japan, and China, female Japanese eels have been
treated with a regular injection of extracts from ground pituitaries of salmon, carp and
catfish to induce egg maturation ŽYamamoto and Yamauchi, 1974; Yamauchi et al.,
1976; Yu et al., 1993.. Final maturation is achieved by giving an injection of 17a ,20b-
dihydroxy-4-pregnen-3-one ŽDHP.. The time of spawning can also be manipulated by
changing the time of DHP administration ŽKagawa et al., 1997.. In the European eel, a
single injection of HCG is sufficient to induce spermatogenesis ŽKhan et al., 1987.. In
practice, one or two HCG injections will ensure the quality and quantity of sperm
released. In the wild, the development of gonads is blocked if reproductive migration
does not occur ŽDufour, 1994.. Maturation could be blocked due to the very small
amount of gonadotropins contained in the pituitary of the adult and the inhibition of
gonadotropin by dopamine ŽDufour et al., 1983, 1988..
Egg quality is related to the nutritional condition of broodstock, and induction
method Žmaterials and dosage.. Broodstock are primarily fed on artificial diets, which
may or may not supply sufficient nutrients for further embryonic and early larval
development. Furthermore, treated eels, suffering from handling stress usually do not
feed, which may affect egg quality. At TFRI, female Japanese eels Ž Ang. japonica. and
European eels Ž Ang. anguilla. were induced to ovulate by the injection of ground catfish
pituitary and HCG for 10 weeks. Females that were ready to be induced had average
weights around 800–1000 g. Regular injection of ground catfish pituitary was conducted
weekly or biweekly. When the body weight increased to over 110%, due to ripe eggs
absorbing water, another injection of ground catfish pituitary was given 48 h after the
regular injection. Then, after 24 h or when egg diameter was 0.9–1.0 mm, 2 mgrkg BW
of DHP was injected. Most of the females spawned within 13–20 h after DHP injection.
It should be noted that the use of 17a-hydroxyprogesterone instead of DHP could lower
the cost of hormone treatment 100 times. Spawning generally occurred around 3 to 6
AM. For convenience, ovulation can also be regulated to daytime by manipulating the
induction time. However, the majority of the eggs are overripe or immature with
multiple oil droplets. Fertilized eggs have a diameter of 0.9–1.0 mm, which later
expands to 1.1 mm after absorbing water. Hatching occurs after 37 h of incubation at
22–23 8C. Eggs of poor quality do not hatch. Larvae of poor quality die soon after
hatching. Aeration bubbles and oil films on the water surface often result in high
mortality of hatched larvae.
Pond-reared male Japanese eels mature more easily than females. Male eels, usually
less than 300 g, are regularly injected with 1–2 IU HCGrg BW every 1–2 weeks. Milt
can be produced after being injected for 5 weeks. Inducing the male eel to produce
sperm is much easier than inducing the female eel to produce eggs.
Larval rearing for the eel is still a mystery. In the past decade, Japanese eel larvae
could be kept for 31 days in Taiwan, 27 days in Japan, and 22 days in China ŽChen,
1993; Yu and Tsai, 1994; Tanaka, 1998.. Recent progress in Japan has prolonged the
survival of Japanese eel larvae for as long as 253 days ŽTanaka, 1999a,b.. For the
European eel, the best reported survival duration is 3.5 days ŽProkhorchik, 1986..
Due to the different ecological habits of eel, rotifers seem to be a less appropriate
food during the preleptocephalus stage. Recent studies have found some organic
materials, such as feces of zooplankton, tubes of tail worms, and suspended organic
particles Žso-called Amarine snowB . in the intestine of eel larvae, implying that the
24
Table 5
Important characters in larviculture of five representative finfish discussed
Characters Kinds of fish
I.C. Liao et al.r Aquaculture 200 (2001) 1–31

Freshwater finfish Milkfish Groupers Cobia Eels
Reason for discussion Ø Freshwater culture Ø Marine culture Ø Marine culture Ø Sea cage aquaculture Ø Challenge of the 21st
Ø Currently popular Ø Easy to propagate Ø Difficult to propagate century
Broodstock Ø Controllable Ø Fairly controllable Ø Difficult Ø Easy Ø Uncontrollable
Induction of maturation Ø Easy Ø Spontaneous Ø Easy Ø Easy Ø Uncertain
Ø Spontaneous
Induction of spawning Ø Spontaneous Ø Spontaneous Ø Hormone management Ø Spontaneous Ø Hormone management
Ø Spontaneous
Larval food Ø Simple Ø Simple Ø Complicated Ø Fairly complicated Ø Very complicated
Larval rearing Ø Easy Ø Easy Ø Difficult Ø Easy Ø Extremely difficult
Limiting factor Ø Inbreeding Ø Low price Ø Egg quality Ø Cannibalism Ø Unknown
Ø Cannibalism Ø Space
Ø Viral infection
Rate of survival to Ø ) 30% Ø 30–50% Ø 3–5% Ø 5–10% Ø 0%
grow-out stage
Rearing period Ždays. Ø 25–35 Ø 20–25 Ø 50–60 Ø 40–50 Ø ) 250
larvae might live on small or dissolved organic materials ŽOtake et al., 1993.. In Japan,
larvae that survived to 27 days were given artificial feed made from ground shark egg
ŽTanaka, 1998.. Larvae that survived more than 250 days were given ground shark egg
mixed with other nutritious materials under a light–dark controlled environment with
intensive bottom-cleaning ŽTanaka, 1999a,b.. This has been a revolutionary break-
through in the larviculture of eel, but the larvae that survived were generally small in
size, the 253-day-old larvae having the largest total length of only 30 mm. In Taiwan,
larvae that survived 31 days were reared in water from a grow-out pond in which certain
unknown factors might be present and beneficial for larval survival ŽYu and Tsai, 1994..
The difference in the morphology, particularly in the shape of the head and teeth,
between hatchery-reared 31-day-old larvae and wild-caught larvae remains a mystery
ŽLiao, 1998; Liao et al., 1999.. The high mortality in larval rearing may be attributed to
nutritional deficiencies andror unsuitable environmental conditions.
The important characteristics of the representative finfish discussed in this paper are
summarized in Table 5.
6. Problems and prospects
Techniques for finfish larviculture in Taiwan have been developed for more than 40
years. Modern larviculture techniques have been applied to many species. However,
problems concerning inbreeding and overproduction still exist. Strategies for disease
control and cannibalism prevention still need to be improved.
The larviculture industry in Taiwan is operated by many small family-scale sub-busi-
nesses. In other countries where aquaculture businesses are organized by large compa-
nies, larviculture operations from broodstock management to larval rearing or even
grow-out are all under one business umbrella. The different sub-businesses in Taiwan,
however, operate independent of each other. The primary concern of each sub-business
is just their own profits and what happens to the other sub-businesses is hardly
considered. In other words, the owners of the different sub-businesses are actually
missing the whole picture. They do not realize that each sub-business affects the other
sub-businesses as well. For instance, communication among fry producers, and between
fry producers and grow-out farmers is very poor. Overproduction that leads to a serious
decline in market price is a problem due to such poor communication. Prices of some
marine finfish, such as milkfish and red snapper, have sharply declined due to
overproduction. Another problem originating from this business structure is the use of
broodstock from local farms, which can cause inbreeding. Dwarfism, low disease
resistance, and abnormality of swimbladder, spine, operculum, and fins in the offspring
are major problems relating to inbreeding.
There is a need to establish an association for regional hatchery operators to share
information and control fry production. More specifically, hatchery operators in one
locality can be organized as a team and teamwork can be promoted among the members.
Local markets should be organized and supervised through a market awareness program
to watch over the market price and prevent overproduction. Furthermore, Taiwanese
products must be strongly promoted and introduced to the international market to avoid
the overproduction problem. To make Taiwan’s hatchery-produced fry internationally

competitive, the following strategies should be implemented: Ž1. a fry quality guarantee
system; Ž2. fry brands or trademarks; and Ž3. market identification or creation. Taiwan
must prepare itself for the global trends of the future.
Disease control of larviculture in Taiwan is also under improvement. The control of
protozoan infestation Žespecially Amyloodinium ocellatum. is still under research and
development. Grouper fry after metamorphosis or at later developmental stages are
sensitive to the white spot disease, caused by Cryptocaryon irritans, especially after the
grading processes. Recently, viral diseases caused by NNV and iridovirus-like virus
have become serious problems. Prevention and control of viral infection are possible
through selection of virus-free broodstock by polymerase chain reaction ŽPCR., stabi-
lization of culture environment, and improvement of nutrition. To prevent the spread of
diseases, a strict quarantine system should be adopted. For instance, PCR can be
incorporated into the quarantine system to monitor viral infection. In addition, hatchery
operators have to be educated on disease prevention techniques. Hatcheries should be
designed to allow rotation of the use of facilities. A hatchery should have at least two
sets of production modules that can be used alternatively. While one module is used for
production, the other can be simultaneously cleaned, disinfected, and prepared for the
next run.
Cannibalism is another serious problem in many species of finfish propagated in the
hatchery. Some studies have already been done and cannibalism is generally controlled
by physical manipulations such as grading and feeding adjustments. However, these
techniques are labor-intensive and sometimes result in physical injuries. Therefore, the
genetic and physiological mechanism of cannibalism must be understood in order to
develop a more effective way of control.
In general, effective approaches are necessary to ensure successful finfish larvicul-
ture. Food is one of the most critical factors that determine larval survival. For every
culture species, the appropriate type of live food containing the nutrients required by the
larvae must be selected. The food must also be given at the right time. Feeding time
should coincide with larval feeding behavior. Furthermore, the food should be provided
in adequate quantity to avoid starvation or water pollution due to overfeeding.
Techniques in larviculture require continuous modification and improvement. New
methods can be introduced to increase the production level. For instance, super-intensive
recirculating culture systems can be developed for larviculture. In such systems,
contamination can be prevented and stocking densities can be increased. Additionally,
the potential of deep ocean water application to finfish larviculture should be explored.
Deep ocean water is free from contamination and rich in dissolved inorganic nutrients.
Although its temperature is low, it can easily be adjusted by heating to the desired
rearing temperature.
Development of larviculture technologies for other species with high aquaculture
potential must be continuously pursued. For instance, people have tried hard to breed
bluefin tuna ŽThunnus thynnus . in captivity. Very recently, the trials have become
successful. Bluefin tuna, kept captive in Tokyo Sea Life Park, spontaneously spawned
successively in the aquarium, which was a cylindrical water tank 30 m in diameter ŽThe
Nihon Keizai Shimbun, Aug. 17, 1999.. Fertilized eggs were obtained and hatched.
Based on this experience, aquaria may be one choice besides cages for breeding large
pelagic marine finfish, especially with respect to the risk of losing fish.
7. Conclusion
The importance of aquaculture will become more apparent in the coming century
with the demanding pressure from an ever-increasing world population. The assurance
of mature and sound techniques in larviculture is the most important channel towards a
sustainable aquaculture industry. Therefore, some strategies should be kept in mind and
carefully carried out. The first is to speed up the completion of unfinished current
research. The second is to formulate and implement systematic regulations and self-con-
trols. An integrated regulation system must be imposed to ensure that the environment is
not harmed by practices of larviculture techniques, that natural traits of propagated fish
are not lost in the process, and that fry are not wasted due to overproduction. Thirdly, it
is important to strengthen international communication and information exchange.
Exchange of technical know-how is necessary to prevent research duplication, which can
waste human resources, to avoid overproduction, and to keep diseases from spreading.
Development of technologies will be speeded up smoothly as international cooperation
is promoted. Lastly, it is time for us to strive for a better earth. When technologies are
developed, nature should be protected as well. Let the people of the future enjoy a better
environment than people of the present and the past.
Acknowledgements
We thank the organizers of the Workshop on Advanced Biotechnology in Finfish

Hatchery, particularly Dr. Cheng-Sheng Lee, for their generous travel and accommoda-
tion supports and warm hospitality to I.C. Liao. We acknowledge the assistance of Mr.
Su-Lean Chang in collecting documents, and Ms. Josephine N. Nocillado and Dr.
Chang-Tai Shih in preparing and reading the manuscript. Thanks are also owed to the
anonymous reviewers of this manuscript.
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Techniques in Finfish Larviculture in Taiwan

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Aquaculture 200 Ž2001.

Techniques in finfish larviculture in Taiwan

0044-8486r01r$ - see front matter q 2001 Published by Elsevier Science B.V.

developments and advanced techniques of finfish larviculture in Taiwan. q 2001 Published by

Keywords: Techniques; Finfish; Larviculture; Taiwan

2.1. Collection of broodstock

A successful hatchery operation is based on healthy broodstock. Broodstock are

To assure good quality eggs, dietary lipid composition in broodstock feed is

2.3. Sexual maturation and spawning

Spawning of broodstock could occur artificially through hormonal induction or

Artificial propagation is further assured with gamete cryopreservation. In Taiwan,

2.4. Egg collection and incubation

3.2. Nutrient requirement

3.4. Pond management

4.2. Oyster trochophores

The following uses cases of several typical aquaculture species to demonstrate

5.1. Freshwater finfish

5.2. Milkfish (Cha. chanos)

5.4. Cobia (R. canadum)

Cobia, also known as sergeant fish, is a middle-sized tropical marine fish. It is

I.C. Liao et al.r Aquaculture 200 (2001) 1–31

6. Problems and prospects

the overproduction problem. To make Taiwan’s hatchery-produced fry internationally

We thank the organizers of the Workshop on Advanced Biotechnology in Finfish

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