Aquaculture 234 (2004) 485 – 496

www.elsevier.com/locate/aqua-online

Effects of Na/K ratio in seawater on growth and

energy budget of juvenile Litopenaeus vannamei
Changbo Zhu, Shuanglin Dong *, Fang Wang, Guoqiang Huang
Mariculture Research Laboratory, Fisheries College, Ocean University of Qingdao,
Qingdao 266003, People’s Republic of China
Received 3 July 2003; received in revised form 27 October 2003; accepted 24 November 2003

Abstract

The effects of seawater Na/K ratio on growth and energy budget of juvenile Litopenaeus
vannamei Boone, 1931 were investigated. Salinity (S = 30 ppt) and the total concentration of sodium
and potassium in the experimental water were kept constant. Seven treatments were set: R1, R2, R3c,
R4, R5, R6 and R7, the Na/K ratios were 25.6, 34.1, 47.3, 85.2, 119.3, 153.3 and 187.4 (mmol/
mmol), respectively. The shrimp in R7 (Na/K = 187.4) died within 2 weeks after the experiment
began. After the 30-day feeding trial, the molting frequency (MF) and feed intake in terms of wet
weight (FIw) and energy content (FIe) of the shrimp under the other six treatments were not
significantly affected by the different Na/K ratios in the seawater. The final wet weight, weight gain
(WG), specific growth rates (SGRw and SGRe) and food conversion efficiencies (FCEd and FCEe) of
the shrimp under Treatment R6 were significantly lower than those under the other five treatments.
There was a trend that, with all the indices mentioned above, the values of the test shrimp under
Treatment R2 was the highest, and the sequence was: R2 (Na/K = 34.1)>R3c (Na/K = 47.3)>R1 (Na/
K = 25.6)>R4 (Na/K = 85.2)>R5 (Na/K = 119.3)>R6 (Na/K = 153.3). In the experiment, the
percentages of energy deposited for growth ( G), energy lost for respiration (R), energy lost in
excretion (U) and energy lost in exuviae (E) to the energy consumed in food (C) were significantly
affected by the different Na/K ratios in seawater, while the percentage of energy lost in feces ( F) to C
was not significantly affected by the ratios. The percentage of G to C of the test shrimp under
Treatment R6 was significantly lower while the percentage of E to C was significantly higher than
those under the other treatments. There were no significant differences among the energy allocations
of the test shrimp under Treatment R1, R2, R3c, R4 and R5. According to regression analysis,

* Corresponding author. Tel.: +86-532-2032827; fax: +86-532-2032799.

E-mail address: dongsl@mail.ouc.edu.cn (S. Dong).

0044-8486/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2003.11.027
486 C. Zhu et al. / Aquaculture 234 (2004) 485–496

optimal shrimp growth could be obtained by regulating the Na/K ratio of the seawater to 40 – 43 at a
salinity of 30 ppt.
D 2004 Elsevier B.V. All rights reserved.

Keywords: Na/K ratio; Seawater; Litopenaeus vannamei; Growth; Energy budget

1. Introduction

The great tolerance to wide salinity range of some penaeid shrimps, such as Penaeus
monodon and Litopenaeus vannamei (Boone, 1931), and the high incidence of epidemic
diseases in coastal areas has led to the growth of inland shrimp farming (Boyd, 2001;
McIntosh and Fitzsimmons, 2001; Jiang and Gong, 2002). Inland shrimp farming was
initiated in Thailand more than a decade ago; it is now undertaken in the United States,
Ecuador, Brazil, China and several other countries. Possibly 30– 40% of P. monodon
culture in Thailand is in inland ponds (Boyd, 2002). By the end of 2002, this activity was
present in nine inland Chinese provinces and the culture area in China has amounted to
14,000 ha (Dong, in press).
Saline water for inland shrimp farming can be obtained in two ways. In some areas,
aquifers containing naturally saline water exist, and ponds can be filled from wells
developed in these aquifers. Where saline water is not available naturally, brine solutions
from coastal salt farms or solid salt may be transported to the ponds and mixed with
freshwater to provide enough salinity for shrimp culture (Boyd, 2001). In most areas, the
latter technique is being much more widely cased by shrimp farmers, which may bring all
out significant short-term economic benefits, but at the risk of creating long-term
cumulative environmental impacts, including excessive freshwater expenditure, and the
salinization of agricultural land and freshwater resources (Flaherty and Vandergeest, 1998;
Fegan, 1999; Braaten and Flaherty, 2000, 2001; Jiang and Gong, 2002). There are no such
problems in culturing shrimp in the saline ground waters in the inland or coastal low-lying
saline-alkaline areas. China has been exploiting and ameliorating the saline-alkaline
wetland by using the fish pond-agricultural terrace system in the Yellow River Delta for
decades. It is now found that this type of aquaculture could not only reduce the salinization
of the soil around the pond area, but also be a good way for the sustainable development of
the agricultural economy (Cheng, 1993; Gu, 1994). However, well waters with adequate
salinity may be unacceptable for shrimp culture because of low concentrations of
potassium, magnesium, or other ions (Boyd, 2002). In the saline-alkaline area of Yellow
River Delta in China and some places of New South Wales, Australia, the saline ground
waters are both chloride type, and most of their chemical compositions are similar to that
of oceanic seawater of the same salinity except potassium in which is 90 – 95% less than
similar salinity oceanic seawater (Wang et al., 2001; Allan and Fielder, 2002). Both places
have the experience of adding potash to fortify the potassium concentration for shrimp or
fish culture (Liu, 2001; Fielder et al., 2001; Allan and Fielder, 2002). It has been reported
that the survival of red drum Sciaenops ocellatus was significantly correlated with the Na+/
K+ and K+/Cl
ratios of the saline ground water (Forsberg et al., 1996). Yet there are few
 C. Zhu et al. / Aquaculture 234 (2004) 485–496 487

reports on the effects of water Na/K ratio on the growth and other physiological characters
of aquatic animals. Therefore, in order to exploit these saline lands, investigating
adaptability to water Na/K ratios is a most pressing matter. The aim of the present
research was to investigate the relation between the growth and energy budget of L.
vannamei and water Na/K ratios, and then consider if Na/K ratio could be used as a water
quality index for shrimp ponds in potassium deficient saline areas.

2. Materials and methods

2.1. Source of juvenile L. vannamei and preliminary acclimation

Juvenile L. vannamei were obtained from the Jiaozhou Shrimp Farm in Qingdao,
People’s Republic of China. When the shrimp were transported to the laboratory, they
were stored in three continuously aerated 600-l fiberglass tanks with natural seawater (29 –
31 ppt) to undergo a 10-day preliminary acclimation to the indoor laboratory conditions,
during which they were fed ad libitum twice a day (8:00 and 18:00) with commercial
shrimp ration pellets (composition: 41.58% crude protein, 8.36% crude lipid, 10.75% ash
and 8.74% moisture; energy content: 19.37 kJ/g dry matter).

2.2. Experimental design and artificial seawater preparation

In order to eliminate the interference of imbalance from other ions, the experimental
water was prepared by adding instant artificial seasalts into fully aerated tap water. The
instant artificial seasalts were specially designed and produced by General Sea Salt
Factory, Ocean University of Qingdao, in which the sodium and potassium ingredients
were precisely compounded so as to keep their total concentration constant while the Na/K
ratios varied. The salinity of the newly prepared artificial seawaters was 30 ppt, and pH
8.2 F 0.1. The total concentration of monovalent cations and other ions were kept
approximately constant. According to a preliminary acute toxicity test, L. vannamei could
just survive in the artificial seawater (S = 30 ppt) within a Na/K ratio range of 25 –204
(mmol/mmol), with a 96-h survival rate above 80%. Therefore, seven treatments were set:
R1, R2, R3c, R4, R5, R6 and R7, and the Na/K ratios were 25.6, 34.1, 47.3, 85.2, 119.3,
153.3 and 187.4 (mmol/mmol), respectively. The Na/K ratio of R3c was identical with the
value of oceanic seawater, and it was set as control. The concentrations of Na+ and K+
were determined with an inductively coupled plasma-atomic emission spectrophotometer
(ICP-OES; VISTA-MPX, VARIAN). The details are given in Table 1.

Table 1
Concentrations of Na+, K+(mmol l
1) and Na/K ratios of the experimental artificial seawater (S = 30 ppt)
Treatments R1 R2 R3c R4 R5 R6 R7
(Na/K ratio) 25.6 34.1 47.3 85.2 119.3 153.3 187.4
Na+ 393.2 398.0 402.0 406.8 408.5 409.5 410.1
K+ 15.4 11.7 8.5 4.8 3.4 2.7 2.2
Na+ + K+ 408.6 409.7 410.5 411.5 411.9 412.1 412.3
488 C. Zhu et al. / Aquaculture 234 (2004) 485–496

2.3. Experimental procedure and acclimation of shrimp

At the end of the 10-day preliminary acclimation, 224 juveniles of similar size
were selected and transferred into glass aquaria (45  25  30 cm) filled with 30
l artificial seawater (S = 30 ppt) of different Na/K ratio for a 7-day acclimation. Seven
treatments, four replicates, in total 28 aquaria were used, each aquarium held eight
shrimp. The aquaria were randomly located. In order to prevent the shrimp from
jumping out, every aquarium was covered with a mesh cover. The ambient temper-
ature was controlled with an air-conditioner. Aeration was provided continuously and
two-thirds of the water volume was exchanged every second day to ensure high water
quality. During acclimation and the following experiment, dissolved oxygen was
maintained above 6.0 mg/l, pH 8.1 F 0.2, water temperature at 25 F 0.5 jC, and a
simulated natural photoperiod (14 light/10 dark) was used. The shrimp were fed ad
libitum twice a day (6:00 and 16:00) with the commercial shrimp ration pellets men-
tioned above.
After the 7-day acclimation and 24 h starvation, the shrimp under each treatment
were weighted individually. To remove excess moisture, shrimp were blotted dry with
paper towel and weighted to the nearest 0.001g using an electronic balance. Then, 22
individuals of similar weight (5.360 F 0.030 g) under each treatment (totally 154) were
selected, of which six were sampled and dried in an oven at 65 jC to constant
weight, homogenized, and stored at
20 jC to estimate the body composition and
energy content of the initial shrimp. The remaining 16 individuals were test shrimp,
which were randomly assigned to four aquaria (4 individuals/aquarium) for the fol-
lowing experiment. The rearing conditions were similar to those during the acclimation
period.

2.4. Samples collection and analysis

During the course of the experiment, the daily food supplied was precisely weighed
and recorded. The uneaten feed and feces were collected by siphon within 3 h after each
meal. Exuviae (molted exoskeletons) were collected and recorded at all times. The
collected uneaten feed, feces and exuviae were dried at 65 jC and kept for further
analysis. The experiment lasted for 30 days, from October 11th to November 11th of
2002. At the end of the experiment, all the test shrimp were collected and dried at 65 jC
for 48 h.

2.5. The estimation of energy budget

The energy content of the shrimp bodies, feed and feces was measured by Parr
1281 Oxygen Bomb Calorimeter. The energy budget was calculated using the
following equation for the crustacean energy budget (Petrusewicz and Macfadyen,
1970):

C ¼GþF þU þEþR
 C. Zhu et al. / Aquaculture 234 (2004) 485–496 489

where, C is the energy consumed in food; G, the energy deposited for growth; F, the
energy lost in feces, U, the energy lost in excretion; E, the energy lost in exuviae,
and R, the energy for respiration.
The estimation of U was based on the nitrogen budget equation (Levine and Sulkin,
1979):

U ¼ ðCN
GN
FN
EN Þ  24 830

where CN is the nitrogen consumed from food; FN , the nitrogen lost in feces; GN, the
nitrogen deposited in shrimp body; EN, the nitrogen lost in exuviae; 24 830, the energy
content in excreted nitrogen per gram (J/g). The nitrogen contents in shrimp bodies, feed,
feces and exuviae were determined by Kjeldahl method.
The value of R was calculated as the following energy budget equation:

R¼C
G
F
U
E

2.6. Calculation and data analysis

Weight gain (WG) and molting frequency (MF) were calculated as follows:

WGð%Þ ¼ 100ðW2
W1 Þ=W1

MF ¼ ð%day
1 Þ ¼ 100Nm =ðNs  T Þ

specific growth rate (SGRw), feed intake (FIw) in terms of the wet weight were calculated
as:

SGRw ð%day
1 Þ ¼ 100ðlnW2
lnW1 Þ=T

FIw ðwt:% body weight day
1 Þ ¼ 100C=½T ðW2 þ W1 Þ=2

food conversion efficiency (FCEd) in terms of dry matter was calculated as:

FCEd ð%Þ ¼ 100ðW 2V
W 1V Þ=C

where, W2 and W1 are the final and initial wet body weight of the shrimp; Nm, the
number of molts; Ns, the number of shrimp; T, the duration of the experiment; C, the
total food consumed; W2V and W1V are the final and initial dried body weights of the
shrimp.
SGR, FI and FCE in terms of energy content (SGRe, FIe, FCEe) were calculated
similarly.
Statistics were performed using SPSS 10.0 statistical software (SPSS, 1999). All data
were subjected to one-way ANOVA. If significant differences were indicated at the 0.05
level, then Duncan’s multiple range tests were used to test the differences between
treatments.
490 C. Zhu et al. / Aquaculture 234 (2004) 485–496

3. Results

3.1. Survival and molting

The survival and molting data of the test shrimp are presented in Table 2. The shrimp
under Treatment R7 (Na/K = 187.4) displayed anorexia, low-activity. Death began after 7
days, and all were dead in 16 days, indicating that L. vannamei could only endure the high
Na/K ratio seawater for a short time (Treatment R7 was excluded in the following analyses
unless specified). The survival of test shrimp under the other six treatments varied from
81.3% to 100%, and mortalities arose from shrimp jumping out when exchanging the
water and cannibalism. There were no statistically significant differences in survival
among those treatments though survival under Treatment R4 was the highest.
According to the result of one-way ANOVA, there were no significant differences
among the molting frequencies (MF) of the test shrimp under all of the six treatments
( p>0.05).

3.2. Growth

At the beginning of the feeding trial, the body weights of the test shrimp under each
treatment were similar (Table 2). At the end of the experiment, the final wet weight of the
test shrimp under Treatment R6 was significantly lower than those under treatments with
lower Na/K ratios. The test shrimp also showed significant differences in weight gains
(WG), in which the test shrimp under Treatment R2 had the highest value and was
significantly higher than those under Treatment R5 and R6; and the value under Treatment
R6 was significantly lower than those under the other treatments.
During the experiment, significant differences also occurred in specific growth rates in
terms of wet weight (SGRw) and energy content (SGRe) among treatments. Similar to the
results of WG, the highest SGRs occurred under Treatment R2, and the values under
Treatment R6 were significantly lower than those under the other treatments. Meanwhile,

Table 2
Growth, survival and molting of L. vannamei in the artificial seawater of different Na/K ratios (mean F SE)*
Treatmentsy Body wet weight (g) WG (%) Survival (%) MF (%
day
1)
Initial Final
R1 5.309 F 0.094 8.212 F 0.113b 54.78 F 2.72bc 81.3 F 6.3 7.29 F 0.71
R2 5.237 F 0.048 8.765 F 0.261b 67.34 F 4.58c 87.5 F 7.2 8.75 F 0.54
R3c 5.439 F 0.068 8.446 F 0.205b 55.44 F 5.29bc 93.8 F 6.3 7.50 F 0.76
R4 5.364 F 0.049 8.262 F 0.214b 54.08 F 4.17bc 100.0 F 0.0 8.75 F 0.54
R5 5.400 F 0.100 7.999 F 0.218b 48.15 F 3.31b 87.5 F 7.2 8.81 F 0.83
R6 5.413 F 0.064 7.188 F 0.365a 32.67 F 5.74a 87.5 F 12.5 10.07 F 0.35
R7 5.360 F 0.030 – – 0 –
WG: weight gain; MF: molting frequency.
* Values (expressed as mean F SE, n = 4) with different letters in the same column are significantly different
from each other ( p < 0.05).
y
The Na/K ratios of treatments R1, R2, R3c, R4, R5, R6 and R7 were 25.6, 34.1, 47.3, 85.2, 119.3, 153.3 and
187.4 (mmol/mmol), respectively.
 C. Zhu et al. / Aquaculture 234 (2004) 485–496 491

there were no significant differences among the test shrimp under Treatment R1, R2, R3c,
R4 and R5 in SGRe, but SGRw under Treatment R5 was significantly lower than that under
Treatment R2 (Fig. 1).

3.3. Feed intake

Feed intakes in terms of wet weight (FIw) and energy content (FIe) are presented in Fig.
2. It is apparent that FIs under Treatment R2 were the highest, and the values under
Treatment R6 were the lowest. However, one-way ANOVA showed that there were no
significant differences in FIs among treatments ( p>0.05), which indicated that the Na/K
ratios of the artificial seawater had no significant effects on the feed intake of L. vannamei
in the experiment.

3.4. Food conversion efficiency

Food conversion efficiencies in terms of dry matter (FCEd) and energy content (FCEe)
manifested the same tendency: FCE under Treatment R2 was the highest, and the value
under Treatment R6 was significantly lower than those under the other treatments, while
there were no significant differences in FCEs among the test shrimp under Treatments R1,
R2, R3c, R4 and R5 (Fig. 3).

3.5. Energy allocation

Table 3 shows the allocation of the energy consumed in food (C) of the test shrimp in
artificial seawater of different Na/K ratios. As compiled, except for the energy lost in

Fig. 1. Specific growth rates (SGR) of L. vannamei during the 30-day experiment. R1, R2, R3c, R4, R5 and R6
exhibit the Na/K ratio of 25.6, 34.1, 47.3, 85.2, 119.3 and 153.3, respectively. Means (n = 4) with different letters
within the same group are significantly different ( p < 0.05), and bars represent standard errors of the means. The
indices SGRw and SGRe indicate specific growth rate in terms of wet weight and energy content, respectively.
492 C. Zhu et al. / Aquaculture 234 (2004) 485–496

Fig. 2. Feed intakes (FI) of L. vannamei during the 30-day experiment. See legend of Fig. 1 for R1 to R6. Bars
represent standard errors of the means (n = 4). The indices FIw and FIe indicate feed intake in terms of wet weight
and energy content, respectively.

feces ( F), the other parts of energy allocation of the test shrimp presented significant
differences among treatments. The test shrimp under Treatment R6 spent much more
energy of the consumed food on respiration and excretion while deposited less energy for

Fig. 3. Food conversion efficiencies (FCE) of L. vannamei during the 30-day experiment. See legend of Fig. 1 for
R1 to R6. Means (n = 4) with different letters within the same group are significantly different ( p < 0.05), and bars
represent standard errors of the means. The indices FCEd and FCEe indicate food conversion efficiency in terms
of dry matter and energy, respectively.
 C. Zhu et al. / Aquaculture 234 (2004) 485–496 493

Table 3
Energy allocation of L. vannamei during the 30-day experiment (mean F SE)*
Treatmentsy G (% C
1) R (% C
1) F (% C
1) U (% C
1) E z (% C
1)
R1 14.58 F 0.53b 62.22 F 1.06b 16.00 F 0.90 5.83 F 0.10b 1.36 F 0.15b
R2 15.57 F 0.45b 61.73 F 1.53b 15.28 F 1.57 5.90 F 0.20b 1.52 F 0.07b
R3c 14.75 F 0.33b 64.42 F 0.50ab 13.28 F 0.43 6.06 F 0.05ab 1.49 F 0.19b
R4 14.04 F 0.76b 64.77 F 0.48ab 13.35 F 0.57 6.11 F 0.07ab 1.72 F 0.09b
R5 13.48 F 0.94b 62.34 F 1.33b 16.41 F 0.89 5.88 F 0.18b 1.89 F 0.17b
R6 7.60 F 2.17a 67.21 F 1.25a 16.04 F 0.94 6.53 F 0.22a 2.62 F 0.26a
* Values (expressed as mean F SE, n = 4) with different letters in the same column are significantly different
from each other ( p < 0.05).
y
The Na/K ratios of treatments R1, R2, R3c, R4, R5 and R6 were 25.6, 34.1, 47.3, 85.2, 119.3 and 153.3
(mmol/mmol), respectively.
z
C is the energy consumed in food; G, the energy deposited for growth; R, the energy for respiration; F, the
energy lost in feces; U, the energy lost in excretion; and E, the energy lost in exuviate.

growth than those shrimp under any other Na/K ratio. In reverse, the shrimp under
Treatment R2 deposited the highest percentage of energy for growth and spent less
energy on respiration and excretion. Meanwhile, the test shrimp under Treatment R6
spent much more energy in exuviae than the shrimp under any other treatment. However,
no significant differences were found in energy allocation among treatments R1, R2, R3c,
R4 and R5 ( p>0.05).

4. Discussion

Generally, L. vannamei can survive within a salinity range of 2 –78 ppt, and can live
normally within a salinity range of 5 –40 ppt (Wang, 2000). According to the present
experiment, improper Na/K ratio might be an important limiting factor for the survival
and growth of L. vannamei. It could only grow normally in saline water within a limited
Na/K ratio range. Fielder et al. (2001) suggested that because salinity of the groundwater
could vary and the interactions between ions during animal osmoregulation were
complex, it might be more important to consider the ratio of ions rather than the
specific concentration of individual ions in the water. In the seven treatments of the
present experiment, the absolute concentrations of K+ changed a lot when the Na/K ratio
varied. Regardless of the Na/K ratios, the absolute concentrations of K+ of the seven
treatments equaled the K+ concentrations of salinities of 54.4, 41.3, 30.0, 16.9, 12.0, 9.5
and 7.8 ppt oceanic seawater. Moreover, it was reported that the growth rates of L.
vannamei juveniles (initial mean weight = 1.6 g) at salinity 5 and 15 ppt were
significantly greater than at 25 and 35 ppt (Bray et al., 1994), seemingly low
K+concentrations (at salinity 5 and 15 ppt) were better for the shrimp. However, the
shrimp could hardly survive under Treatment R7, and the growth rates were much better
under Treatment R2 and R3c than those under R4, R5 and R6. It is difficult to explain
such phenomena merely in terms of K+ concentrations. Similarly, there was no effect of
salinity (3 –6 ppt) on survival or growth of red drum S. ocellatus, but its survival in
groundwater (S = 15 ppt) was negatively correlated with the ratio of Na+/K+ ( p = 0.02,
494 C. Zhu et al. / Aquaculture 234 (2004) 485–496

r2 = 0.60) and positively correlated with the ratio of K+/Cl
( p = 0.05, r2 = 0.50)
(Forsberg et al., 1996).
In the present research, different Na/K ratios in the artificial seawater significantly
affected the growth, feed intake and energy allocation of juvenile L. vannamei. The final
wet weight and weight gain of the test shrimp under Treatment R2 were both much greater
than under any other treatment. Also, R2 had the highest SGRs (even higher than those in
the control). Through regression analysis, the correlation of SGRw and Na/K ratio (X) of
the artificial seawater was obtained from a non-linear (quadratic) curve:

SGRw ¼ 1:4577 þ 0:0043X
5  10
5 X 2 ðp < 0:001; r2 ¼ 0:567Þ:

The maximum value occurred while X = 43. Similarly, the relationship between weight
gain (WG) and Na/K ratio (X) of the artificial seawater was

WG ¼ 56:1792 þ 0:1611X
0:002X 2 ðp < 0:001; r2 ¼ 0:547Þ;

and the maximum value occurred while X = 40.3. Accordingly, juvenile L. vannamei could
get the best growth within Na/K ratios of 40– 43 under a salinity of 30 ppt. Such values are
lower than the Na/K ratio of oceanic seawater namely 47.3. It might therefore be more
favorable for the growth of juvenile L. vannamei if the water K+ concentration was
comparatively higher than that of oceanic seawater of the same salinity. Therefore, to some
extent, higher K+ concentration or lower Na/K ratio could promote the growth and food
conversion efficiency of juvenile L. vannamei. However, when the K+ concentration is too
high or Na/K ratio is too low, contrary results occurred. In Treatment R7 of the present
experiment where K+ was seriously deficient, the test shrimp displayed anorexia, low-
activity and death. The best growth obtained in Treatment R2 where K+ concentration was
higher than that in the oceanic seawater. Similar results were reported in the snapper
Pagrus auratus cultured in K+ deficient saline groundwater in New South Wales, Australia
(Fielder et al., 2001).
No significant differences were demonstrated in feed intake among treatments in the
test shrimp, thus the growth of juvenile L. vannamei was affected by Na/K ratios through
FCE and the percentage of energy deposited for growth ( G) to the energy consumed in
food (C) (see Fig. 3 and Table 3). In the present experiment, the shrimp under Treatment
R2 had the highest percentage of G to C, and showed the best growth. On the contrary, the
shrimp under Treatment R6 showed the lowest percentage of G to C; as a result the growth
was the worst. It was remarkable that the percentage of exuviae (E) to C in the shrimp
under Treatment R6 was significantly higher than those under other treatments, which
coincided with its high molting frequency (see Table 2). In fact, it was because the Na/K
ratio of the artificial seawater in Treatment R6 was so different from the value of the
oceanic seawater that stress occurred in the test shrimp. Many factors could enhance
molting but have adverse effects on the growth of shrimp (Vijayan and Diwan, 1995), such
as low salinity to P. monodon (Allan and Maguire, 1992) and ammonia to Penaeus
japonicus (Chen and Kou, 1992). High molting frequency could not only increase the
energy expenditure for exuviae, but also affect the entire energy allocation strategy of the
animal, for the process of molting involves consumption of oxygen and energy (Water-
 C. Zhu et al. / Aquaculture 234 (2004) 485–496 495

man, 1960; Penkoff and Thurberg, 1982; Cockcroft and Wooldridge, 1985). In conclusion,
it was feasible to take Na/K ratio as an index for considering the suitability of potassium
deficient saline groundwater for shrimp culture and for the management of water quality.
However, because of the complexity and variation of the groundwater chemical compo-
sitions, there are still many problems to overcome for exploiting the aquaculture potential
of natural saline groundwater.

Acknowledgements

This work was funded by the Project under the Major State Basic Research of China
(Grant No. G1999012011), the National 10th 5-Year Major Program (Grant No.
2001BA505B) and the State Agriculture Program (Grant No. K2002-16). We thank
General Sea Salt Factory, Ocean University of Qingdao, for designing and producing the
experimental instant seasalts.

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