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ABSTRACT
In this article, we present an overview of milkfish (Chanos chanos) culture, in order to provide
basic information on its life history, reproductive biology, fry production, culture methods, and genetics
as well as its commercial importance. Milkfish is a warm-water marine species, and its culture is
widespread in the Asian-Pacific region; it is a unique species in the family Chanidae. Currently
milkfish is an important brackish water aquaculture species in Southeast Asia and represents an
important component of the fisheries sector and national economy in Indonesia, the Philippines,
and Taiwan; in Taiwan, annual total production has surpassed 450,000 metric tons since 2000.
However, milkfish production is faced with limitations, the most significant of which is the unpredic-
table and limited supply of fry. Milkfish production comes mostly from aquaculture, and the availability
of fry for stocking plays an important role in the success and development of the industry. Traditionally
milkfish production relied on the capture of fry from the wild; but important attempts have been made
to develop milkfish broodstock-hatchery technology, and significant successes have been achieved
in the artificial propagation of this species.
Key words: Milkfish, Chanos chanos, Life history, Reproductive biology, Culture methods.
1
Department of Tropical Agriculture and International Cooperation, National Pingtung University of Science and
Technology, Pingtung, Taiwan 912, R.O.C.
2
Department of Aquaculture, National Pingtung University of Science and Technology, Pingtung, Taiwan 912,
R.O.C.
* Corresponding author. E-mail: mctseng@mail.npust.edu.tw
230 Franklin S. Martinez, Mei-Chen Tseng and Sin-Ping Yeh
Fresh water
Juvenile (5)
Sub adult
(6) Fry (4)
Adult stage (7) Larvae
(3)
Lagoon, lakes,
swamps
Yolk sac larvae Mangroves
Spawning Embryo (1) (2) swamps, estuaries
ground in the
Ocean
Fig. 1. Life history of milkfish represented by seven developmental stages from embryo to adults (modified from
Lee, 1995).
Taiwan
Hawaii
J F M A M J J A S O N D
Fig. 2. Spawning season (months) of different milkfish population (Lee, 1995; Tucker, 1998).
benthic algal filter-feeders. At this point, limits reported to be from 15 to 40°C, but the
juveniles are found in large quantities in coastal ideal temperature is between 20 and 33°C
estuaries, mangrove areas, coastal lagoons, with an optimum for spawning of 28°C (Landau,
and swamps and begin moving upriver into 1992; Lee, 1995). Others factors such as
lakes. Then juveniles develop into subadults currents and wind may also influence the
at different sizes (200~4500 g) and at different selection of spawning areas in order to ensure
ages (1~4 years). Most subadult fish leave the survival of eggs and larvae (Lee, 1995).
the inshore environment and return to the The natural spawning seasons differ
ocean where they survive on a diet of plankton according to milkfish populations (Fig. 2); the
and eventually spawn; but other milkfish reproductive status can be determined
remain in fresh water (Lee, 1995). based on different criteria; but histological
Milkfish spawn in clear oceanic waters, examination of gonadal development is
usually at less than 40 m in depth over sand considered the most accurate method of
or coral about 30 km offshore during the determining spawning seasons (Lee, 1995).
warm months of the year (Landau, 1992). Milkfish are able to spawn more than once
Despite this species being able to tolerate a during the annual spawning season (Lee et
wide range of environmental conditions, there al., 1997). Prijono et al. (1988) demonstrated
are many factors which affect milkfish's that spawning occurred on nine occasions
selection of spawning grounds; temperature within 4 months after stocking in a group of
is one of the most important factors, with wild brood stock (specimens with a weight
Milkfish Culture in the Asian-Pacific Region 231
Weight FCR
140 2
c
120
b
1.5
100 a
Weight (mg)
80
FCR
a 1
60
b
40 c 0.5
20
0 0
0 16 34
Salinity (ppt)
Fig. 3. Effects of salinity on growth and the feed conversion rate (FCR) of milkfish fry. Different letters indicate
a significant difference at p < 0.05 (modified from Alava 1998).
232 Franklin S. Martinez, Mei-Chen Tseng and Sin-Ping Yeh
Table 1. Milkfish, aquacultures methods, and production in Indonesia, Taiwan, and the Philippines
(FitzGerald, 2004)
Country Culture method Stocking density Production
(fish/ha) (kg/ha/year)
Indonesia Extensive (shallow-water) 6000 300~1000
Taiwan Extensive 6000~7000 1800~2500
(shallow-water)
Semi-intensive > 25,000 8000~12,000
(deep-water)
Philippines Extensive 1500~6000 800~4000
Pen 30,000~40,000 2000~10,000
Cage 35~100 fish/m3 13~17 kg/m3
Note that the production varies within and between these countries, which can be explained by differences in
production efficiency in each country.
Milkfish Culture in the Asian-Pacific Region 233
portable nets which surround groups of fry characteristics such as the presence of three
and conduct them into a small area. minute opening pores in the urogenital region
in females, while the males only have two
Fry production through induced matura- main external openings (Garcia, 1988; Lee,
tion and spawning of brood stock 1995).
Once milkfish have maturated, in order to
Even though artificial propagation has accurately determine the time for spawning,
progressed in the last two decades, a source it is better to induce spawning in females by
of wild fry is still required in some locations the administration of exogenous hormones
(Leis and Reader, 1991; Lee et al., 1997). (fish pituitary gonadotropins, HCGs, and LHRH-
Nowadays the production of fry is possible a). Hormone therapy can be administered
due to the capture of subadults from the wild through food, pellet implants, and intramus-
and their rearing until they become sexually cular and intraperitoneal injections. Different
mature. Adults naturally mature during the dosages have been studied; but 200 µg/kg
spawning season when they reach 3 kg or using a pellet implant or injection seems to
more at 4 and 5 years old for males and females, be a better dose in terms of fertilized spawn
respectively. Maturation can also be induced and fertilization rate. Fish spawn 24~48 h
through the use of exogenous hormones such after treatment depending on the hormone
as luteinizing hormone-releasing hormone therapy. A mature female can spawn 2
(LHRH-a), fish pituitary gonadotropin (salmon million eggs/kg of body weight per year (Lee,
pituitary homogenate and carp pituitary 1995).
homogenate), human chorionic gonadotro- Lee et al. (1997) and FitzGerald (2004)
pins (HCGs), and gonadal steroids (estradiol described two basic hatchery operation
17-β and 17 α- methyltestosterone), as well methods for producing milkfish fry: intensive
as through environmental stimulation, such and semi-intensive. In the intensive system
as temperature and manipulating the photo- larvae are stocked at a relatively high density
period in order to increase the number of of 20~30 larvae/l, and tanks are employed for
daylight hours (Lee, 1995). Milkfish seem to larval rearing with the addition of separately
be a gonochoristic species. Determination of cultured algae, rotifers, and Artemia at
sex is important for the successful breeding in different stages during the culture period (Fig.
captivity; sex can be distinguished by external 4). In the semi-intensive system, a pond is
Maturation of
broodstock
Transfer to grow-out
Spawning phase
(natural or induced)
Enrichment
Fig. 4. Schematic representation of hatchery facilities needed to produce milkfish fry (FitzGerald, 2004).
234 Franklin S. Martinez, Mei-Chen Tseng and Sin-Ping Yeh
utilized, and natural growth of phytoplankton species is best suited for culture in the tropics
and zooplankton is stimulated through fertiliza- because of its fast growth, efficient use of
tion; in this system, larvae are stocked at a natural food, propensity to consume a variety
lower density (5 larvae/l). of supplemental feeds, resistance to diseases
Both systems begin with fertilized eggs and handling, and tolerance to a wide range
(1.1~1.25 mm in diameter). Eggs need to be of environmental conditions (Lim et al., 2002).
incubated in saltwater (30~34 ppt) and a In nature, milkfish are generally herbivorous,
range of temperatures of 26~30°C, and good but in some conditions can be considered
aeration is also required. In the intensive omnivorous. Lückstädt et al. (2001) and
system, eggs are directly stocked in the Lückstädt and Reiti (2002) analyzed the
larval rearing or incubator tanks at a density stomach contents of adult milkfish fed natural
of 1000 eggs/l, and then larvae are trans- food and reported that the stomach contents
ferred to special areas to receive their starter consisted mainly of single-cell green and
diet (exogenous feed). In Taiwan, hatchery blue-green algae, but also contained benthic
operators use the semi-intensive system, and planktonic organisms and crustacean
while the intensive system is used in Hawaii, larvae. In captivity, various types of feed
the Philippines, and Indonesia to produce can be used such as live feed for rearing
milkfish fry (Lee, et al., 1997; FitzGerald, larvae and artificial or formulated feed for fry,
2004). juveniles, and later stages of development
At optimal temperatures, the eyes become (Lee, 1995).
entirely pigmented and the mouth is open at Like other marine fish species, milkfish
54 h after hatching; the yolk sac is almost do not have absolute protein requirements, but
completely reabsorbed 120 h after hatching do require a balanced mixture of essential
(Bagarinao, 1986). Exogenous feed must amino acids (Table 2) (Lim et al., 2002).
be provided before the yolk sac is completely Generally marine fish are not able to convert
absorbed. Rotifers (Branchionus plicatilis) shorter-chain fatty acid (linoleic acid, 18:3 n-3
are considered the standard first feeding and linolenic acid, 18:2 n-6) to longer-chain
regimen at 10~20 rotifers/ml, but also micro- fatty acids, so it is necessary to provide these
algae and Artemia nauplii are used (Bagarinao, through the diet. Lipids and fatty acids play
1986; Lee, 1995). Normally 21-day-old milkfish significant roles due to their function in
fry (14~16 mm in length) are harvested and maintaining cell membrane structures, stress
transferred to nursery ponds for 30~45 days tolerance, and proper development and
of rearing prior to grow-out culturing (Lee, functioning of neural and visual systems
1995; Hilomen-Garcia, 1997). (Borlongan, 1992; Alava, 1998; Faulk and
Holt, 2005). The polyunsaturated fatty acids,
NUTRITION AND FEED specifically eicosapentaenoic acid (EPA, 20:5
n-3), arachidonic acid (ARA, 20:4 n-6), and
In terms of feeding habits, milkfish are docosahexaenoic acid (DHA, 22:6 n-3), have
considered herbivorous (Hertrampf and Piedad- been shown to be essential in the diet of
Pascual, 2000; Lim et al., 2002). This marine fish larvae (Faulk and Holt, 2005).
Table 2. Essential amino acid requirements for juvenile milkfish (Lim et al., 2002)
Amino acid Requirement Amino acid Requirement
(% of dietary protein)1 (% of dietary protein) 1
Arginine 5.2 Methionine 2.5
Histidine 2.0 Phenylalanine 4.2
Isoleucine 4.0 Threonine 4.5
Leucine 5.1 Tryptophan 0.6
Lysine 4.0 Valine 3.6
1
For diets containing 40%~45% protein.
Milkfish Culture in the Asian-Pacific Region 235
Essential fatty acid (EFA) deficiency symptoms herbivorous and artificial diets containing
consist of poor growth, low feed efficiency, 23%~27% crude protein (Lim et al., 2002).
anemia, and high mortality especially in The crude protein contents as well as the
milkfish larvae (Fig. 5). Ascorbic acid is also proportions of other ingredients used to
required in larval fish diets; scoliosis, twisted prepare artificial diets for fry, grow-out, and
gill filaments, and short operculae are some broodstock stages are presented in Table 3.
of the signs of ascorbate deficiency (Gapasin Feed peas (Pisum sativum), an abundant
et al., 1998). agricultural product, is a potential feed
Milkfish accept a variety of diets such as ingredient used in fish diets. It is a high-
meal forms and sinking particles or floating energy, medium-protein ingredient with a
pellets. Using pellets improves the feeding protein content of 22%~24% and 14.3 kJ/g of
efficiency due to the physical characteristics digestible energy (Borlongan et al., 2003). The
such as better stability which prevents them efficiency of feed peas as a feed ingredient
from dissolving and nutrients separating in has been evaluated for milkfish (Borlongan et
the water. The most common pellet size al., 2003) as well as trout (Oreochromis
used for feeding fish at 400 to 500 g is mykiss) (Hertrampf and Piedad-Pascual,
approximately 4~5 mm in diameter and 6~8 2000). However, they can also be used as
mm long (Lim et al., 2002). Newly hatched dietary protein source for other aquaculture
fry use their yolk sac as a source of nutrients; species.
a brief explanation of first feeding was presented Borlongan et al. (2003) found significant
above, so the information provided in this differences in both survival and FCR after 12
part mostly concerns fry, juvenile, and adult weeks of feeding different diets. The highest
stages. survival and lowest FCR were obtained in
Diets for fry usually contain a high milkfish fed the control diet (containing fish
percentage of crude protein, but this is meal, soybean meal, meat and bone meal,
decreased as the fish grow. For example, and copra meal as the principal protein
fry require a diet with 40% crude protein, and sources) as well as D-2 and D-3 (containing
then at the grow-out phase, milkfish are fed 5% and 10% substitution of the total protein
Fig. 5. Percentage of incidence of opercular deformities among 40- day-old milkfish larvae. Control, larvae fed
Chlorella-cultured rotifers and newly hatched Artemia; HUFAs only, larvae fed rotifers and Artemia
enriched with high unsaturated fatty acids (HUFAs); HUFAs + C, larvae fed rotifers and Artemia
enriched with HUFAs + vitamin C. For each trial, different letters for the treatment means indicate a
significant difference (p < 0.05) (modified from Gapasin et al., 1998).
236 Franklin S. Martinez, Mei-Chen Tseng and Sin-Ping Yeh
Table 3. Feed formulae for milkfish in different stages of development (Lim et al., 2002)
Ingredient (%) Fry Grow-out Broodstock
(40% protein) (27% protein) (36% protein)
Fish-meal 30.0 10.0 23.0
Soybean meal 20.0 34.5 33.0
Shrimp meal 15.0
Wheat flour 25.45 15.0
Grains by product 48.0
Marine fish oil 8.0 2.0 2.5
Rice bran 21.0
Pellet binder 3.0 3.0
Vitamin mix 1.0 0.5 1.0
Dicalcium phosphate 1.5 1.0
Antioxidant 0.05
Trace mineral mix 0.5 0.5 0.5
Total 100.0 100.0 100.0
100 3
90 a
c b b
80 d 2.5
e e e
Survival rate (%)
70 2
60
c c c
FCR
50 1.5
40 b b b
30 1
a a
20 0.5
10
0 0
D-1 (0%) D-2 (5%) D-3 (10%) D-4 (15%) D-5 (20%) D-6 (25%)D-7 (30%) CMF
Diet (% of subtitution of total protein with pea meal)
Fig. 6. Response of milkfish juveniles to the various diets after 12 weeks of feeding. The control diet (D-1)
contained fishmeal, soybean meal, meat and bone meal, and copra meal as the principal protein
sources. Feed pea meal was progressively substituted at 0% (D-1), 5% (D-2), 10% (D-3), 15% (D-4),
20% (D-5), 25% (D-6), and 30% (D-7) of the total protein and was compared with commercial milkfish
feed (CMF). Different letters for the treatment means indicate a significant difference at p < 0.05
(modified from Borlongan et al., 2003).
with pea meal, respectively). However, it natural food. Normally feed is offered two or
was observed that milkfish fed diets with up three times daily and is distributed by hand or
to 15% and 20% levels of feed pea meal had automatic feeders according to the biomass
better FCRs than those on the commercial in the ponds or cages. Supplementary feed
feed control. (Fig. 6) can be offered at a rate (% of biomass) of
As with other aquaculture species, the 20% for fry and 9% for juveniles, and at 3%~
feeding rate depends on the fish size and % of body weight for growth to harvest size
environmental conditions like water tempera- (Lim et al., 2002).
ture, dissolved oxygen, and the presence of
Milkfish Culture in the Asian-Pacific Region 237
Table 4. Principal disease problems in milkfish under aquaculture conditions (Lee, 1995)
Type of Description Biological causes Physical factors Treatment and
disease prevention
Bacterial
infections
Red spot or - Red spot and ulcers on Vibrio anguillarum - Cold water - Vaccination
vibriosis the body (mainly on the Prevention: temperatures - Bacteriophage
abdominal region) Lower stocking density - Overcrowding - Antibiotics
- Hemorrhage of eyes, and provision of good - Poor water - Disinfectant bath
mouth, and inner water quality quality
surface of gills
- Pale gills
Fin rot or - Frayed fins Flexibacter columnaris - Changes in pH - Antibiotics
columnaris - Hemorrhagic wounds Prevention: and temperature, (oxytetracycline,
disease - Excess of white mucus Lower stocking density Low dissolved sulphadiazine)
on the skin and provision of good oxygen - Malachite green
- Missing scales water quality, plus - High ammonia - Copper sulfate
- Lesions in the mouth avoidance of folic acid and levels
inositol
Septicemia - Loss of scales Aeromonas hydrophila - Stressed fish - Antibiotic therapy
- Hemorrhages of the Prevention: under poor water (oxytetracycline)
gills, anus, mouth, and Lower stocking density conditions
fins and provision of good
- Skin ulcers water quality
- Accumulation of ascetic
fluid
Mycotic
Infections
Milky eye - White adipose lid over Fungal infection - Stress after Bath using:
infection the eyes handling - Potassium
- Fungal hyphae in Prevention: (sampling, permanganate
lesions Reduce fish handling as transport, - Malachite green
much as possible harvesting)
- Low water
temperature
Parasitic
Infections
Copepoda - Loss of weight Genus Lernae - Overcrowding Bath using:
- Gill damage Prevention: - Warm water - 3%~5% NaCl
- Inflammation and ulcers Drain and dry the bottom temperature - Sodium chloride
at the Point of of ponds
attachment
strated that the histidine concentration as the fasting period increased from 0 to 60
decreased by approximately 46% after 60 days, and the hepatosomatic index dramatically
days of starvation, compared to the milkfish decreased after the first 10 days of starvation
control group, which were fed normally during (Fig. 7). Additionally starvation resulted in
the same period of time. The author also important reductions in proteins and fat of
demonstrated that the body weight decreased 11% and 54%, respectively, in white muscle
Milkfish Culture in the Asian-Pacific Region 239
Weight HIS
50 2
a
40 a
ab 1.5
bc
Weight (g)
30 b c
HSI (%)
c d 1
20 d
d
0.5
10
0 0
0 10 25 40 60
Period of starvation (days)
Fig. 7. Reduction in body weight and the hepatosomatic index (HIS) of milkfish during 60 days of starvation
(modified from Shiau et al., 2001).
19 a a b
c 1.5
ab
18
bc 1
17 c 0.5
16 0
0 10 25 40 60
Period of starvation (days)
Fig. 8. Reduction in the protein and crude fat composition in white muscle of milkfish during 60 days of starva-
tion (modified from Shiau et al., 2001).
of milkfish (Fig. 8). the ponds. Juario (1988) and Pillay & Kutty
Special care should be taken during milk- (2005) described the harvest methodology
fish manipulation due to consumer preferences, related to the culture system.
particularly when fish is marketable in a fresh - Completely draining the ponds is a method
condition. Milkfish harvesting techniques that very few farmers use because fish
depend on the rearing method and design of harvested in this manner may have a
240 Franklin S. Martinez, Mei-Chen Tseng and Sin-Ping Yeh
300
250
Thousands metric tons
200
150
100
50
B
450
400
Value (thousands US$)
350
300
250
200
150
100
50
0
1990 1992 1994 1996 1998 2000 2002 2004
Years
Fig. 9. Major milkfish producing countries: Indonesia, the Philippines, and Taiwan. (A) aquaculture production
and (B) values from 1990 to 2004 (FAO FishStat database, 2004).
Milkfish Culture in the Asian-Pacific Region 241
Prijono, A., I. Tridjoko, A. Nyoman Adiasmara Giri, Nations, Regional Office for Asia and the
W. E. Poernomo, C. Vanstone, T. Daulay. Pacific, Bangkok. 52 p.
(1988). Natural spawning and larval rearing of Sumagaysay-Chavoso, N. S., and San Diego-
milkfish in captivity in Indonesia. Aquacul- McGlone, M. L. (2003). Water quality and
ture, 74: 127-130. holding capacity of intensive and semi-inten-
Seng, L. T. and Colorni, A. (2002). Infectious sive milkfish (Chanos chanos) ponds. Aquacul-
diseases of warmwater fish in marine and ture, 219: 413-429.
brackish water In Diseases and Disorders of Taiwan Fisheries Year Book. (2004). Retrieved
Finfish in Cage Culture, Woo, P. T. K., D. W. January 06, 2005 from: http://www.fa.gov.tw/
Bruno, L. H. S. Lim, Cabi Publishing. New chn/statistics_price/year_book/
York, USA, 193-230. Tucker, W. J. (1998). Marine Fish Culture. 1st
Shiau, C. Y., Y. J. Pong, T. K. Chiou and Y. Y. Tin. edition. Kluwer, Harbor Branch Oceano-
(2001). Effect of starvation on free histidine graphic Institution and Florida Institute of
and amino acids in white muscle of milkfish Technology, Melbourne, FL, USA. 750 p.
Chanos chanos. Comparative Biochemistry Villaluz, A. C. (1988). Milkfish (Chanos chanos
and Physiology Part B, 128: 501-506. Forskal) collection and handling. Southeast
Sugiyama, S., D. Staples and S. Funge-Smith. Asian Fisheries Development Center, Philippines.
(2004). Status and potential of fisheries and Retrieved January 06, 2005 from: http://www.
aquaculture in Asia and the Pacific. Food fao.org/docrep/field/003/AC282E/AC282E04.
and Agriculture Organization of The United htm
244 Franklin S. Martinez, Mei-Chen Tseng and Sin-Ping Yeh
虱目魚養殖之現狀與趨勢
Franklin S. Martinez1, Mei-Chen Tseng2* and Sin-Ping Yeh2
在這篇文章㆗,我們綜述現今虱目魚(Chanos chanos)的生物㈻研究,提供其生態生活
史,繁殖㈻,仔魚產量,養殖方法,遺傳㈻㆖的㈾訊和商業㆖之重要性。虱目魚是虱目魚
科㆗唯㆒的魚種,屬暖㈬性海㈬魚類,在亞洲-太平洋區域被廣泛的養殖,現在是東南亞㆞
區最重要的養殖魚種。對於㊞尼、菲律賓和台灣而言,虱目魚養殖佔㈲重要的經濟價值。
㉂ 2000 年開始,台灣每年總產量超過㆕㈩㈤萬公噸。然而,虱目魚產量面臨極限,最主要
的原因是仔魚供給㆖的限制。傳統㆖虱目魚產量是依靠野生魚苗的捕獲,但是虱目魚的種
魚㆟工孵化場,已經成功的完成了㆟工繁殖。目前虱目魚產量最主要來㉂於㆟工養殖,在
台灣成功和發達的㈽業㆗,虱目魚㆟工繁殖魚苗的外銷,在經濟㆖佔㈲重要的㆞位。
關鍵詞:虱目魚,生活史,生殖生物㈻,養殖方法。
1
國立屏東科技大學熱帶農業研究所
2
國立屏東科技大學水產養殖系
* 通訊作者