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Combined effects of water exchange regimes and

Aquaculture Research, 2006, 37, 664^670

doi:10.1111/j.1365-2109.2006.01478.x

Combined effects of water exchange regimes and
calcium carbonate additions on growth and survival of
hatchery-reared juvenile spotted babylon (Babylonia

areolata Link 1807) in recirculating grow-out system
S Kritsanapuntu1, N Chaitanawisuti2,W Santhaweesuk2 & S Y Natsukari3
1

Faculty of Technology and Management, Prince of Songkla University, Suratani,Thailand

2

Aquatic Resources Research Institute, Chulalongkorn University, Bangkok,Thailand

3

Faculty of Fisheries, Nagasaki University, Nagasaki, Japan


Correspondence: N Chaitanawisuti, Aquatic Resources Research Institute, Chulalongkorn University, Bangkok 10330,Thailand. E-mail:
nilnajc1@hotmail.com

Abstract

Introduction

To determine a suitable culture environment to maximize growth and survival, the hatchery-reared juvenile spotted babylon, Babylonia areolata, were held in
plastic rearing tanks at four calcium carbonate additions of 0,100 and 300 g tonne À1, and four water exchange regimes of 0-,15-,30- and 60-day intervals in
a recirculating grow-out system for120 days. The results clearly showed that growth was greatest between water exchange regimes of 15- and 30-day intervals and all calcium carbonate additions, with
water exchange regimes of 0- and 60-day intervals
resulting in poor growth. Final survival was highest
between water exchange regimes of 15- and 30-day
intervals, and all calcium carbonate additions, with
water exchange regimes of 0-day intervals and all
calcium carbonate additions resulting in high mortalities. This study showed that water exchange regimes had a stronger in£uence on the growth of
juvenile B. areolata than calcium carbonate additions. It is recommended that B. areolata juveniles be
maintained within the water exchange regimes
range of 15^30-day intervals and at calcium carbonate additions between 0 and 500 g tonne À1, providing optimum conditions for production of this species
in a recirculating grow-out system.

Although large-scale rearing of Babylonia areolata
in Thailand was technically feasible using £owthrough systems in concrete/canvass ponds, the disadvantages of these systems that must be solved during grow-out of spotted babylon are: these systems
generally require large quantities of water; location
of production systems must be near the sea; stock is
vulnerable to external water supply and quality problems; and growth rate is signi¢cantly in£uenced by
water £ow (Chaitanawisuti, Kritsanapuntu & Nutsukari 2002, 2004). A recirculating grow-out system
had been used for growing ¢sh and shell¢sh for more
than three decades. This system may o¡er an alternative to pond aquaculture technology of this species.
The interest in recirculating grow-out systems was
due to their perceived advantages, including greatly
reduced land and water requirements, high degree
of environmental control allowing productive cycle
growth at optimum rates, feasibility of locating in
far proximity from the sea, and a major issue of water
conservation and reuse (Lazur & Britt 1997; Losordo,
Masser & Rakocy 1998; Masser, Rakocy & Losordo
1999). Research on recirculating grow-out systems
can provide major leaps in spotted babylon culture
intensi¢cation, technology and the understanding of
water quality management for maximium pro¢t by
increasing production, lowering costs and conserving water. Calcium is a very important element for
¢sh and shell¢sh because it is necessary for a variety

Keywords: Babylonia areolata, recirculating growout system, calcium carbonate, water exchange,
growth, survival

664

r 2006 The Authors. Journal Compilation r 2006 Blackwell Publishing Ltd


Aquaculture Research, 2006, 37, 664^670

Combined e¡ects of water exchange regimes S Kritsanapuntu et al.

of functions such as bone and scale growth, muscle
contraction, transmission of nerve impulses, hormone
secretion, intracellular signalling, against osmotic and
ionic grains and losses, and against most environmental toxicants (Calta 2000). Calcium is required in
rather high levels by aquatic animals in comparison
with other mineral elements. These requirements are
met by dietary resources; however, dissolved calcium
is readily taken up by gills of ¢sh/shell¢sh (Robinson,
LaBomascus, Brown & Linton 1987). However, little is
known about the e¡ects of water exchange regimes
and calcium carbonate on growth and survival for
growing of juvenile spotted babylon to marketable
sizes in recirculating systems in Thailand. The aim of
this study was to determine the combined e¡ects of
calcium carbonate additions and water exchange regimes on the growth and survival of juvenile B. areolata in a recirculating grow-out system.
Materials and methods
Experimental recirculating grow-out system
Each plastic culture tank of 50 L capacity was an independent recirculating system with an air lift pump and
a biological ¢lter tank. The bottom area of the culture
tank was 0.78 m2. The biological ¢lter tank of 20 L capacity contained shell fragments and gravels as ¢lters.
Water £owed from the culture tank through the ¢lter
via air lift pumps at a £ow rate of 200 L h À1 before it
was returned to the culture tank. As in the £owthrough system, the tank bottom was covered with a
5 cm layer of coarse sand (0.5^1.0 mean grain size) to
serve as a substrate. Water depth was 30 cm. When
water exchange was done for each treatment, the substrate was cleaned by £ushing it with a jet of water and
sun dried for 6 h. Thereafter, the tanks were re¢lled
with new ambient natural seawater. Filters were rinsed
in seawater to remove particulate matter, sun dried for
6 h, and returned to the ¢lter tanks. Temperature was
controlled to within room temperature Æ 1.5 1C. Salinity was monitored daily, as necessary, to keep the variation within Æ 2.0 ppt by addition of fresh water to
correct for any increased salinity because of water evaporation. Each culture tank had no aeration and photoperiod was naturally 12L:12D. Water quality of pH,
dissolved oxygen, alkalinity, nitrite-nitrogen and ammonia-nitrogen were analysed before water exchanges.
Experimental animals
Juvenile spotted babylons, B. areolata, used in growth
and survival experiments were produced from a

private hatchery, Rayong province, located on the
eastern coast of the Gulf of Thailand. Individuals
from the same cohort were sorted by size to prevent
possible growth retardation of small spotted babylons
when cultured with larger ones. Initial shell length
and whole body weight of spotted babylon averaged


1.12 Æ 0.13 cm (0.98^1.23 cm) and 0.38 Æ 0.16 g
(0.26^0.54 g), n 5 300 respectively. Each experiment
had mean shell lengths and body weights that were
not statistically di¡erent and this allowed for treatments to be compared statistically. Juveniles were
held in experimental culture tanks as described
above at optimal densities of 300 individuals m À 2 or
250 snails tank À1 (Chaitanawisuti & Kritsanapuntu
1999).

Experimental design
The study was conducted from February to May
2004 at Spotted Babylon Aquaculture Research and
Training Unit, Chulalongkorn University, Petchaburi,
Thailand. This laboratory experiment was designed
to test the combined e¡ects of four water exchange
regimes (0-, 15-, 30- and 60-day intervals) and four
levels of calcium carbonate additions (0, 100, 250
and 500 g tonne À1) on the growth and survival for
juveniles of B. areolata. The experiment was 4 Â 4
factorial design, with all 16 water exchange and calcium carbonate combinations test. For water exchange regimes, complete seawater was exchanged
at each designed time interval, and re¢lled with new
ambient seawater. For calcium carbonate additions,
90% calcium carbonate powder of designed amounts
was dissolved homogeneously in seawater and rinsed
in culture tanks.

Rearing method
The snails were fed ad libitum with fresh trash ¢sh,
once daily, at 09:00 hours. The amount of food consumed was recorded daily and uneaten food was
removed immediately after the animals stopped
feeding, and air dried for a period of 10 min before
weighing. Size grading of snails in all treatments
was not done throughout the grow-out period. No
chemical and antibiotic agent was used throughout
the entire experimental periods. The experiment
was terminated over a 120-day culture period. To determine growth performance, 50% of snails were
randomly sampled from each treatment at 15-day intervals, and shell length and whole body weight were

r 2006 The Authors. Journal Compilation r 2006 Blackwell Publishing Ltd, Aquaculture Research, 37, 664^670

665


Combined e¡ects of water exchange regimes S Kritsanapuntu et al.

determined. Shell length was measured with callipers to the nearest mm from the maximum anterior
to posterior distance of shell, and whole weight was
measured after air drying for a period of 10 min before weighing, and then returned to the tank. The
number of dead individuals was recorded at 15-day
intervals. Average shell length increments, body
weight gains and growth rates were calculated after
the method of Chaitanawisuti and Kritsanapuntu
(1999). Body weight gains (BWf^BWi) and monthly
growth rates for body weight ((BWf^BWi)/T) were calculated, where BWf is the mean ¢nal body weight,
BWi the initial body weight andT the time in months.
Mortality, expressed as the percentage of the initial
stocking density, was calculated from the di¡erence
between the number of snails stocked and harvested.

Statistical analysis
Data were analysed with the SPSS statistical package
(version 10). Two-way analysis of variance (ANOVA)
was used to test the interaction of water exchange regimes and calcium carbonate additions at a 5 0.05,
and di¡erences between means were compared
using Tukey’s test at a 5 0.05.

Aquaculture Research, 2006, 37, 664^670

growth of juvenile B. areolata than calcium carbonate additions. At all calcium carbonate additions,
body weight gain was signi¢cantly greater at water
exchange regimes of 15- and 30-day intervals than
at water exchange regimes of 0- and 60-day intervals, with water exchange of 0- and 60-day intervals
resulting in poor growth (Table 1). On the other hand,
at all water exchange regimes, body weight gain was
signi¢cantly greater at calcium carbonate additions
of 0, 100 and 500 g tonne À1 than at calcium carbonate addition of 250 g tonne À1 (Table1). The monthly
growth rates in body weight of B. areolata in the four
water exchange regimes and four calcium carbonate
additions are presented in Fig. 2. The highest overall
growth rates in body weight (0.42^0.52 g month À 2)
were achieved for juveniles grown at water exchange
regimes of15-day intervals and all calcium carbonate
additions. The lowest overall growth rates in body
weight occurred for juveniles grown at all calcium
carbonate additions and water exchange regimes
of 0- and 60-day intervals (0.26^0.37 g month À 2)
(Tables 2 and 3).

3

Results
Growth in body weight
The growths, expressed as body weight gains, of
B. areolata in the four water exchange regimes and
four calcium carbonate additions are presented in
Fig. 1. Two-way ANOVA performed on growth showed
that the e¡ect of water exchange regimes was statistically signi¢cant (Po0.05), and interaction between
water exchange regime and calcium carbonate addition was found. The results clearly showed that water
exchange regimes had a stronger in£uence on the

Body weight gains (g)

2.5
2
1.5
1

0 days
15 days
30 days
60 days

0.5
0

0

100

250

500

Calcium carbonate addition (g / ton)

Figure 1 Body weight gains of juveniles, Babylonia areolata, in recirculating grow-out system at four water
exchange regimes and four calcium carbonate additions.

Table 1 Average body weight gains (Æ SD) of Babylonia areolata juveniles in recirculating grow-out system with four calcium
carbonate additions and four water exchange regimes

Water exchange
(day interval)
0
15
30
60
Average Æ SD

Calcium carbonate additions (g tonne À 1)
0
1.40
1.74
1.52
1.48
1.54

100
Æ
Æ
Æ
Æ
Æ

0.04
0.18
0.08
0.03
0.15a

1.19
1.68
1.44
1.38
1.42

250
Æ
Æ
Æ
Æ
Æ

0.04
0.50
0.02
0.02
0.20a

1.05
1.82
1.27
1.12
1.32

Average Æ SD

500
Æ
Æ
Æ
Æ
Æ

0.04
0.05
0.02
0.04
0.35ab

1.13
1.68
1.49
1.29
1.39

Æ
Æ
Æ
Æ
Æ

0.01
0.04
0.06
0.03
0.24a

1.19
1.73
1.43
1.32

Æ
Æ
Æ
Æ

0.15a
0.07b
0.11c
0.15ac

Data are means of three replicates. Di¡erent subscriptions indicate signi¢cant di¡erences (Po0.05).

666

r 2006 The Authors. Journal Compilation r 2006 Blackwell Publishing Ltd, Aquaculture Research, 37, 664^670


Aquaculture Research, 2006, 37, 664^670

Combined e¡ects of water exchange regimes S Kritsanapuntu et al.

Growth in shell length
Shell length increments of B. areolata in the four
water exchange regimes and four calcium carbonate
additions are presented in Fig. 3. Two-way ANOVA performed on shell length increments showed that the
e¡ect of water exchange regimes was statistically signi¢cant (Po0.05). The results clearly showed the similar trends as body weight gain. At all calcium

carbonate additions, shell length increment was signi¢cantly greater at water exchange regimes of 15and 30-day intervals than at water exchange regimes
of 0- and 60-day intervals (Table 4). On the other
hand, at all water exchange regimes, shell length increment was signi¢cantly greater at calcium carbonate additions of 0, 100 and 500 g tonne À1 than at
calcium carbonate addition of 250 g tonne À1 (Table
4). The highest overall growth rates in shell length

0.7

2
Shell length increments (cm)

Growth rate (g/mo)

0.6
0.5
0.4
0.3
0 days
15 days
30 days
60 days

0.2
0.1
0

0

100
250
500
Calcium carbonate addition (g / ton)

Figure 2 Growth rates in body weight of juveniles, Babylonia areolata, in recirculating grow-out system at four
water exchange regimes and four calcium carbonate additions.

1.8
1.6
1.4
1.2
1
0.8
0.6

0 days
15 days
30 days
60 days

0.4
0.2
0

0

100
250
500
Calcium carbonate addition (g / ton)

Figure 3 Shell length increments of juveniles, Babylonia
areolata, in recirculating grow-out system at four water
exchange regimes and four calcium carbonate additions.

Table 2 Average growth rates in body weight (Æ SD) of Babylonia areolata juveniles in recirculating growout system with
four calcium carbonate additions and four water exchange regimes

Water exchange
(day interval)
0
15
30
60
Average Æ SD

Calcium carbonate additions (g tonne À 1)
0
0.35
0.52
0.47
0.37
0.43

100
Æ
Æ
Æ
Æ
Æ

0.01
0.05
0.02
0.01
0.08a

0.29
0.42
0.36
0.34
0.35

250
Æ
Æ
Æ
Æ
Æ

0.01
0.02
0.01
0.01
0.05b

0.26
0.46
0.32
0.28
0.33

Average Æ SD

500
Æ
Æ
Æ
Æ
Æ

0.01
0.01
0.01
0.01
0.09b

0.28
0.42
0.35
0.32
0.34

Æ
Æ
Æ
Æ
Æ

0.01
0.01
0.03
0.01
0.06b

0.29
0.46
0.38
0.33

Æ
Æ
Æ
Æ

0.04a
0.05b
0.07ac
0.04a

Data are means of three replicates. Di¡erent subscriptions indicate signi¢cant di¡erences (Po0.05).

Table 3 Average shell length increments ( Æ SD) of Babylonia areolata juveniles in recirculating growout system with four
calcium carbonate additions and four water exchange regimes

Water exchange
(day interval)
0
15
30
60
Average Æ SD

Calcium carbonate additions (g tonne À 1)
0
0.85
1.22
1.04
0.91
1.01

100
Æ
Æ
Æ
Æ
Æ

0.08
0.07
0.08
0.06
0.16a

0.79
1.04
0.90
0.87
0.90

250
Æ
Æ
Æ
Æ
Æ

0.10
0.02
0.05
0.05
0.10b

0.64
0.94
0.87
0.80
0.81

Average Æ SD

500
Æ
Æ
Æ
Æ
Æ

0.05
0.05
0.05
0.11
0.13b

0.90
1.12
1.07
1.10
1.05

Æ
Æ
Æ
Æ
Æ

0.08
0.02
0.13
0.09
0.10a

0.79
1.08
0.97
0.92

Æ
Æ
Æ
Æ

0.11a
0.12b
0.09c
0.13ac

Data are means of three replicates. Di¡erent subscriptions indicate signi¢cant di¡erences (Po0.05).

r 2006 The Authors. Journal Compilation r 2006 Blackwell Publishing Ltd, Aquaculture Research, 37, 664^670

667


Combined e¡ects of water exchange regimes S Kritsanapuntu et al.

Aquaculture Research, 2006, 37, 664^670

Table 4 Average growth rates in shell length (Æ SD) of Babylonia areolata juveniles in recirculating growout system with
four calcium carbonate additions and four water exchange regimes
Calcium carbonate additions (g tonne À 1)

Water exchange
(day interval)

0

0
15
30
60
Average Æ SD

0.21
0.30
0.26
0.22
0.25

100
Æ
Æ
Æ
Æ
Æ

0.02
0.02
0.02
0.03
0.04a

0.22
0.26
0.22
0.21
0.23

250
Æ
Æ
Æ
Æ
Æ

0.03
0.05
0.03
0.03
0.02a

0.15
0.24
0.19
0.20
0.19

Average Æ SD

500
Æ
Æ
Æ
Æ
Æ

0.02
0.02
0.04
0.03
0.04b

0.23
0.28
0.27
0.27
0.26

Æ
Æ
Æ
Æ
Æ

0.03
0.01
0.06
0.01
0.02a

0.20
0.27
0.24
0.23

Æ
Æ
Æ
Æ

0.04a
0.03b
0.04c
0.03ac

Data are means of three replicates. Di¡erent subscriptions indicate signi¢cant di¡erences (Po0.05).

0.4

Growth rate (cm/mo)

0.35
0.3
0.25
0.2
0.15

0 days
15 days
30 days
60 days

0.1
0.05
0

0

100
250
500
Calcium carbonate addition (g/ ton)

Figure 4 Growth rates in shell length of juveniles, Babylonia areolata, in recirculating grow-out system at four
water exchange regimes and four calcium carbonate additions.

(0.19^0.30 cm month À 2) were achieved for juveniles
grown at water exchange regimes of 15- and 30-day
intervals and all calcium carbonate additions (Fig. 4).
The lowest overall growth rates in shell length occurred for juveniles grown at all calcium carbonate
additions and water exchange regimes of 0- and
60-day intervals (0.15^0.27 cm month À 2) (Table 5).

Final survival
The ¢nal survivals of B. areolata in the four water exchange regimes and four calcium carbonate additions are presented in Fig. 5. Two-way ANOVA
performed on ¢nal survival showed that the e¡ect of
water exchange regimes was statistically signi¢cant
(Po0.05), and interaction between water exchange
regime and calcium carbonate addition was found
(Table 5). Tukey’s test showed that at all calcium carbonate additions, ¢nal survival was greater at lower
water exchange of 15- and 30-day intervals than
those at water exchange regimes of 0- and 60-day

668

intervals, and at all water exchange regimes, ¢nal
survival was greater at lower calcium carbonate additions of 0, 100 and 250 g tonne À1 than at calcium
carbonate addition of 500 g tonne À1 (Table 5). The
highest overall ¢nal survivals (65.55^93.52%) were
achieved for juveniles grown at water exchanges of
15- and 30-day intervals and all calcium carbonate
additions. The lowest overall ¢nal survival occurred
for juveniles grown at all calcium carbonate additions and water exchanges of 0-day intervals (47.55^
80.00%) (Table 5).

Discussion
The results of the present study clearly showed that
growth was greatest between water exchange regimes of 15- and 30-day intervals and all calcium
carbonate additions, with water exchange regimes
of 0- and 60-day intervals resulting in poor growth.
Final survival was highest between water exchange
regimes of 15- and 30-day intervals, and all calcium
carbonate additions, with water exchange of 0-day
intervals and all calcium carbonate additions resulting in high mortalities. The results showed that
water exchange regimes had a stronger in£uence on
the growth of juveniles of B. areolata than calcium
carbonate additions. It is recommended that B. areolata juveniles be maintained within the water exchange regimes range of 15^30-day intervals and at
calcium carbonate additions between 0 and
500 g tonne À1 provided optimum conditions for production of this species in a recirculating grow-out
system. Chaitanawisuti and Kritsanapuntu (1999)
reported that the average monthly growth rate of
spotted babylon in a £ow-through culture system in
concrete/canvass tanks was 1.4 g month À1. Food
conversion ratio and ¢nal survival were 1.6% and
95.8% respectively. Chaitanawisuti, Kritsanapuntu

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Aquaculture Research, 2006, 37, 664^670

Combined e¡ects of water exchange regimes S Kritsanapuntu et al.

Table 5 Average ¢nal survivals (Æ SD) of Babylonia areolata juveniles in recirculating growout system with four calcium
carbonate additions and four water exchange regimes
Calcium carbonate additions (g tonne À 1)

Water exchange
(day interval)

0

0
15
30
60
Average Æ SD

60.89
91.58
93.52
92.88
84.72

100
Æ
Æ
Æ
Æ
Æ

0.28
0.22
0.18
0.36
15.91a

80.00
90.44
90.44
82.67
85.89

250
Æ
Æ
Æ
Æ
Æ

0.17
0.23
0.19
0.10
5.37a

73.55
94.44
91.55
66.44
81.49

Average Æ SD

500
Æ
Æ
Æ
Æ
Æ

0.31
0.15
0.18
0.25
13.64a

47.55
73.11
65.55
57.11
60.83

Æ
Æ
Æ
Æ
Æ

0.28
0.21
0.26
0.38
11.00b

65.49
87.39
85.27
74.78

Æ
Æ
Æ
Æ

14.35a
9.67b
13.21b
16.04ab

Data are means of three replicates. Di¡erent subscriptions indicate signi¢cant di¡erences (Po0.05).

140

Final survival (%)

120
100
80
60
0 days
15 days
30 days
60 days

40
20
0

0

100
250
500
Calcium carbonate addition (g/ton)

Figure 5 Final survival of juveniles, Babylonia areolata,
in recirculating grow-out system at four water exchange
regimes and four calcium carbonate additions.

and Santhaweesuk (2004) reported that the average
growth rate of juvenile spotted babylons was
3.86 mm month À1 in length and 1.47 g month À1 in
weight after 8 months when cultured at a density of
300 snails m À 2 in a £ow-through system and
3.21mm month À1 and 1.10 g month À1 when held in
a recirculating system. In recirculating systems,
good water quality must be maintained for maximum growth and for optimum e¡ectiveness of
bacteria in the bio¢lter. A key to successful recirculating production systems is the use of cost-e¡ective
water treatment systems components. All recirculating production systems remove waste solids, oxidize
ammonia and nitrite-nitrogen, remove carbon dioxide, and aerate or oxygenate the water before returning it to the culture tank. Calcium is a very important
element for ¢sh and shell¢sh because it is necessary
for a variety of functions such as bone and scale
growth, muscle contraction, transmission of nerve
impulses, hormone secretion, intracellular signalling, against osmotic and ionic grains and losses,
and against most environmental toxicants. Calcium

enters the ¢sh through the gills, intestines and skin.
The gills are a particularly important calcium uptake
site (Calta 2000). Calcium is required in rather high
levels by aquatic animals in comparison with other
mineral elements. These requirements are met by
dietary resources; however, dissolved calcium is
readily taken up by gills of ¢sh/shell¢sh and some
species can acquire 65^80% of their metabolic needs
from the water (Robinson et al. 1987). Masser et al.
(1999) reported that most recirculating systems are
designed to replace 5^10% of the system volume each
day with new water. This amount of exchange prevents the build-up of nitrates and soluble organic
matter that would eventually cause problems. In
some situations, su⁄cient water may not be available
for these high exchange rates. A complete water exchange should be done after each production cycle
to reduce the build-up of nitrate and dissolved organics. For emergency situations, the recirculating system has an auxiliary water reservoir equal to one
complete water exchange and the reservoir should
be maintained at the proper temperature and water
quality. Hincks and Mackie (1997) reported that maximum growth of zebra mussel (Dreissena polymorpha)
occurred at calcium levels of 32 mg Ca L À 1, an alkalinity of 65 mg CaCO3 L À 1 and a total hardness of
100 mg CaCO3 L À1. There was negative growth at
calcium levels less than 31mg CaCO3 L À1, and positive growth of juveniles zebra mussel only occurred
at pH greater than 8.3. Rakocy (1989) reported that
recirculating systems for ¢sh culture generally recycle 90^99% of the culture water daily and the rearing tank is aerated as in £ow-through systems with
low exchange rates. The recirculating rate (amount
of water exchange per unit of time) can be determined by dividing the volume of water in the tank
by the capacity of the pump. Increasing the number
of turnovers per day would provide increased bio¢ltration, greater nitri¢cation and reduced ammonia

r 2006 The Authors. Journal Compilation r 2006 Blackwell Publishing Ltd, Aquaculture Research, 37, 664^670

669


Combined e¡ects of water exchange regimes S Kritsanapuntu et al.

levels. Most ¢sh production recirculating systems are
designed to provide at least on complete turnover per
hour (24 cycles per day). On the basis of the results of
this study, B. areolata juveniles should be held at
water exchange regimes between 15- and 30-day intervals and calcium carbonate additions between 0
and 500 g tonne À1 in recirculating grow-out systems. Within these culture parameters, juvenile
growth and survival will be optimal, and the e⁄cient
utilization of water and reduction in operation costs
will be maximized.

Acknowledgments
We thank the National Research Council of Thailand
(NRCT), who provided funds for this research in ¢scal
year 1996^2005. We are specially grateful to Associated Professor Dr Padermsak Jarayabhand, Director
of Aquatic Resources Research Institute, Chulalongkorn University, for his encouragement and facilities.

References
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