Aquaculture 306 (2010) 352–356

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Aquaculture
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Effect of salinity on survival, growth, food consumption and haemolymph osmolality

of the pink shrimp Farfantepenaeus subtilis (Pérez-Farfante, 1967)
Emanuell Silva ⁎, Nathalia Calazans, Marcelo Soares, Roberta Soares, Silvio Peixoto
Universidade Federal Rural de Pernambuco, Departamento de Pesca e Aqüicultura, Laboratório de Maricultura Sustentável, 52171-900, Recife, PE, Brazil

a r t i c l e i n f o a b s t r a c t

Article history: The effect of salinity on survival, growth, food consumption and haemolymph osmolality was determined for
Received 7 August 2009 the pink shrimp Farfantepenaeus subtilis in a series of independent experiments. The first experiment, shrimp
Received in revised form 6 January 2010 at PL50 were transferred without salinity acclimation from 30‰ to the salinities of 0, 5, 10, 15, 20, 25, 30 and
Accepted 29 April 2010
35‰ for 96 h. The tolerance of shrimp to abrupt changes in salinity (96 h LS50) was estimated. The second
experiment, growth and survival of shrimp were analyzed during 75 days in the treatments corresponding to
Keywords:
Farfantepenaeus subtilis
the salinities of 5, 15, 25 and 35‰, where the food consumption was measured during four consecutive days.
Salinity Finally, haemolymph osmolality of juveniles (1.23 g ± 0.26) maintained at 5, 10, 15, 20, 25, 30 and 45‰ for
Survival 24 h was evaluated. All PL50 transferred from 30‰ to 0‰ died within 24 h, while survival after 96 h of those
Growth transferred to 5 and 10‰ was 96.6 and 86.6%, respectively. PL50 transferred to salinities higher than 15‰
Food consumption presented no mortality during the test. Based on these results, the lethal salinity (96 h LS50) was estimated at
Haemolymph 12‰. Final weight of shrimp reared at 25‰ was significantly higher than the other treatments, except for
treatment 35‰, which did not differ from shrimp reared in salinity 15‰. Shrimp reared at 5‰ had the lowest
weight. Survival of shrimp varied from 48% (15‰) to 65% (35‰), showing higher values at 25 and 35‰, but
not differing significantly from 5‰. Shrimp reared at 25 and 35‰ had the highest food consumption values.
The isosmotic point estimated from a regression between haemolymph and water osmolality was
377.07 mosM kg− 1, which is equivalent to salinity 14‰. Results indicate that F. subtilis presents higher
growth in salinities higher than its isosmotic point, i.e., at salinities above 25‰.
© 2010 Elsevier B.V. All rights reserved.

1. Introduction (Ferraris et al., 1987). Similarly, in the rearing environment, salinity is

The pink shrimp Farfantepenaeus subtilis is distributed from the
Caribbean Sea (Cuba and Lesser Antilles) to Cabo Frio, Rio de Janeiro,
Brazil (D'Incao, 1995). Previous initiatives to culture this species in
northeast Brazil were constrained mostly by the lack of technology
available and inferior growth performance when compared to the exotic
Litopenaeus vannamei (Nunes et al., 1997). Recently, due to L. vannamei
disease outbreak, the culture potential of F. subtilis has been recon-
sidered as an alternative species for aquaculture in Brazil. However,
there is little information about essential biological and physiological
aspects that will provide a basis for F. subtilis culture.
As most penaeid shrimps, F. subtilis postlarvae are carried by
currents to the estuaries, where they reach the juvenile stage, and
then migrate to the ocean reaching the adult stage (Isaac et al., 1992).
In estuarine environments, penaeids are exposed to abrupt changes in
salinity as a result of tides, winds and rain action (Dalla Via, 1986;
Kumlu and Jones, 1995). Therefore, physiological adaptation to
salinity has a crucial role on the survival of shrimp to adult stage
⁎ Corresponding author. Tel.: + 55 81 3320 6524.
E-mail address: emanuellfelipe@yahoo.com.br (E. Silva).
one of the most important factors, affecting growth, survival and food
consumption of shrimp (Jory, 1995; Kumlu et al., 1999, 2000).
The effects of salinity on growth and survival of several penaeid
species have been reviewed by Dall et al. (1990). Salinity in the range
of 15 to 25‰ is considered ideal for the rearing of Penaeus monodon
(Ferraris et al., 1986a; Chen et al., 1995), while superior growth
performance was observed in the range 20–30‰ for Fenneropenaeus
chinensis (Chen et al., 1995; Zhang et al., 1999). Although the highest
survival of L. vannamei juveniles was found in salinities over 20‰
(Ponce-Palafox et al., 1997), this species has been reared under low
salinity conditions (Laramore et al., 2001; McGraw et al., 2002;
Atwood et al., 2003; Saoud et al., 2003). Food consumption may also
be affected by salinity, as observed for Fenneropenaeus merguiensis
juveniles that showed lower value of consumption in high salinity
(45‰) (Vinod et al., 1996). In contrast, Wasielesky et al. (2003) found
no significant differences in food consumption for Farfantepenaeus
paulensis in salinities between 2 and 36‰.
The level of stress and physiological adaptation of penaeids to
different salinities can be monitored through their osmoregulatory
capacity, which represents the difference between the haemolymph
and medium osmolality (Charmantier et al., 1989; Sang and Fotedar,
2004). Lignot et al. (2000) showed that osmoregulatory capacity

0044-8486/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2010.04.025
 E. Silva et al. / Aquaculture 306 (2010) 352–356
varies for penaeid species according to their size, nutritional condition
and developmental stages.
The present study aimed to determine the effect of salinity on
survival, growth, food consumption and haemolymph osmolality of
the pink shrimp F. subtilis.
2. Materials and methods
2.1. Experimental animals
Ten day-old F. subtilis postlarvae (PL10) were obtained from the
spawning of wild broodstock in a commercial hatchery located in Rio
Grande do Norte, Brazil. These were transferred to our laboratory and
maintained in a fiberglass tank (500 L) under constant aeration,
ambient temperature of 27–28 °C and salinity 30‰. Daily 5% of the
water was renewed. Food consisted of a commercial diet with 40%
crude protein (Purina 40CR2) offered three times daily (0800, 1200
and 1600 h) in the proportion of 20% of total biomass per day, and
Artemia nauplii offered ad libitum once daily (1800 h).
2.2. Tolerance to changes in salinity
To evaluate the tolerance to abrupt changes in salinity, postlarvae
(PL50) were transferred without salinity acclimation from 30‰ to the
treatments corresponding to salinities of 0, 5, 10, 15, 20, 25, 30
(control) and 35‰. The experimental system consisted of 2 L plastic
containers stocked with ten postlarvae each for 96 h with three
replicates per treatment.
The water salinity for each treatment was adjusted one day before
the test by diluting sterilized marine water with tap water, which was
previously aerated for at least 24 h. Salinity was measured using a
handheld refractometer. Experimental conditions were no water
renewal, temperature of 26 °C, photoperiod of 12 h light:12 h dark,
and constant aeration that maintained dissolved oxygen levels above
4 mg/L. Shrimp were fed twice daily (0800 and 1600 h) with the
commercial diet described above in the proportion of 10% of shrimp
biomass/day.
Survival was recorded at 24, 48, 72 and 96 h of exposure to each
salinity level. Lack of response to mechanical stimuli was the criteria
to determine death. Dead shrimp were removed at the time of each
observation. Lethal salinity (LS50) was determined using the software
Trimmed version 1.5 (Hamilton et al., 1977).
2.3. Growth, survival and food consumption at different salinities
Before the experiment, thirty postlarvae were randomly sampled,
blotted on absorbent paper and weighed using an analytical balance
(0.0001 g) to estimate the initial weight. The effect of salinity on
growth, survival and food consumption was analyzed in the
treatments corresponding to the salinities of 5, 15, 25 and 35‰,
with three replicates each.
Shrimp were gradually acclimated from salinity 30‰ to the
respective salinity level during 96 h. Groups of 20 postlarvae were
then stocked in 40 L experimental tanks, which received recirculated
water (24 L/h) from an individual biological filter consisting of 2 L
plastic containers filled with ceramic rings substrate. Controlled
environmental conditions included temperature of 26 °C, 12 h
light:12 h dark photoperiod and constant aeration. Temperature, pH
and dissolved oxygen were measured daily using a multiparameter
(YSI 556). The water in each tank was renewed 50% every week. The
experiment lasted 75 days.
Two commercial pelleted feeds were used. The first one had 40% of
crude protein (Purina 40CR2) and was offered to shrimp for the first
fifteen days at a rate of 10% of total biomass, and another with 35%
crude protein (Purina C35) at a rate of 20% of biomass. Feed was
offered twice daily (0800 and 1600). Shrimp survival and body wet
353
weight were recorded every two weeks by counting and weighing ten
individuals from each tank (30 per treatment), respectively. Shrimp
were blotted dry in tissue paper prior to weighing in an analytical
balance (0.01 g).
On day 30, food consumption was estimated for four consecutive
days. A known amount of feed (10% of shrimp biomass) was offered in
trays. Residual feed were collected after 6 h, placed in pre-weighed
filters and oven-dried (60 °C) until a constant weight. Estimates of
food consumption were adapted from Soares et al. (2005):
FC ð%Þ = f½ðF0 −Fr Þ × Fl  = TBg⁎100
where: FC (%) = food consumption (% of biomass/ meal), F0 = dry
weight of feed offered (g); Fr = dried weight of feed recovered (g); Fl =
difference of feed lost by leaching (%); TB: total shrimp biomass in the
tank.
In addition, three tanks without shrimps were used to estimate the
amount of feed lost by leaching and recovery failure.
2.4. Haemolymph osmolality
F. subtilis juveniles (1.23 g ± 0.26) were transferred without
previous acclimation from salinity 30‰ to 5, 10, 15, 20, 25, 30 and
45‰. Five liter plastic containers were stocked with five shrimp each
according to the different salinity treatments. Water temperature was
maintained at 28 °C. After 24 h, shrimp were blotted dry in tissue
paper and haemolymph samples were taken individually. Only
shrimp in the intermolt stage were used. The haemolymph was
drawn from the pericardial cavity between the cephalothorax and
first abdominal segment using a 1.0 ml syringe. Samples were placed
in Eppendorf tubes and a 10 µl subsample was drawn with a
micropipette to osmolality reading. In addition, the medium osmo-
lality was measured in three water samples from each salinity
treatment using a Wescor® vapor pressure osmometer, model 5520.
The isosmotic point was calculated based on the linear regression
analysis between haemolymph and medium osmolality.
2.5. Data analysis
Data on growth, survival, food consumption and haemolymph
osmolality were analyzed by one-way analysis of variance (ANOVA),
followed by Duncan's Multiple Range Test to determine differences
among treatments.
3. Results
3.1. Tolerance to changes in salinity
Survival of F. subtilis postlarvae to abrupt changes in salinity was
severely affected only when they were submitted to salinity 0‰. In
this case, 100% mortality was observed after 24 h. In the salinities of 5
and 10‰, shrimp survival after 96 h was 96.6 and 86.6%, respectively.
In contrast, for all other salinity treatments (15 to 35‰) survival after
96 h was not affected. The lethal salinity (LS50) of F. subtilis (PL50) for
96 h was estimated at 12‰.
3.2. Growth, survival and food consumption at different salinities
Mean (±SD) water temperature, pH and dissolved oxygen were
26.6 ± 0.6 °C, 8.01 ± 0.55 and 5.42 ± 0.63 mg/L, respectively. From day
45 onwards, the mean final weight of shrimp reared in 25‰ was
significantly higher than those reared at 5 and 15‰ (Table 1). At the
end of the experiment (75 days), the weight of shrimp in treatment
25‰ was significantly higher than the other treatments, except for
treatment 35‰, which did not differ from shrimp reared in salinity
354 E. Silva et al. / Aquaculture 306 (2010) 352–356

Table 1 wide tolerance to abrupt changes in salinity. Lethal salinity (LS50)

Mean (± SE) values of weight (g), survival (%) and food consumption (% of biomass/ varies widely among penaeids and depends on several factors such as
meal) of shrimp F. subtilis reared at different salinities for 75 days.
temperature and life stage of the individuals (Brito et al., 2000). Olin
Variables Day Salinity (‰) and Fast (1992) observed an increase in salinity tolerance of P.
5 15 25 35 monodon and L. vannamei until PL20 and PL15, respectively. F. paulensis
a a a increased its resistance to changes in salinity during development
Weight 0 0.06 ± 0.01 0.06 ± 0.01 0.06 ± 0.01 0.06 ± 0.01a
(n = 10) 15 0.14 ± 0.01ab 0.13 ± 0.01ab 0.15 ± 0.01a 0.11±0.01b from PL15 to PL30 (Tsuzuki and Cavalli, 2000). Charmantier-Daures et
30 0.23 ± 0.01b 0.26 ± 0.02ab 0.29 ± 0.02a 0.25 ± 0.01ab al. (1988) observed that the LS50 decreased in Marsupenaeus japonicus
45 0.38 ± 0.02b 0.39 ± 0.03b 0.51±0.03a 0.46 ± 0.02ab PL20–60 with increasing temperature, but a similar pattern was not
60 0.76 ± 0.03b 0.82±0.05b 0.97 ± 0.04a 0.87 ± 0.04ab found for F. chinensis. In M. japonicus, the LS50 for 24 h decreased from
75 1.01 ± 0.05c 1.12 ± 0.07bc 1.38 ± 0.06a 1.27 ± 0.08ab
Survival 0 100a 100a 100a 100a
larval stages to postlarvae (PL10) (Charmantier-Daures et al., 1988).
15 95 ± 0a 98.3 ± 1.0a 96.6 ± 1.0a 98.3 ± 1.0a The LS50 for F. subtilis PL50 in this study was estimated at 12‰, which
30 85 ± 1.8a 90 ± 0ab 95 ± 0bc 96.6 ± 1.0c is slightly higher than the LS50 around the salinity 10‰ found for PL28
45 78.3 ± 3.8ab 68.3 ± 1.0a 81.6 ± 4.5ab 85 ± 3.1b of Farfantepenaeus brasiliensis (Brito et al., 2000) and F. chinensis
60 70 ± 3.6ab 55 ± 3.1b 73.3 ± 2.7a 71.6 ± 4.2a
(Charmantier-Daures et al. 1988). Although the LS50 values estimated
75 61.1 ± 5.8a 48.3 ± 2.1b 56.6 ± 4.2ab 65 ± 4.8a
Food 30–34 2.93 ± 0.16b 3.20 ± 0.12ab 3.38 ± 0.15a 3.46 ± 0.13a here suggest that F. subtilis postlarvae may be less resistant to lower
consumption salinities, it agrees well with LS50 values reported for penaeids at
Different superscript letters in the same row indicate significant differences (p b 0.05).
similar life stages (Dall et al., 1990).
In the juvenile stage of F. subtilis, growth performance and survival
recorded in this study were improved between 25 and 35‰, which is
15‰. Shrimp reared at 5‰ had the lowest weight, which differed in accordance with results from penaeids such as F. chinensis,
significantly from treatments 25 and 35‰. Farfantepenaeus californiensis, F. brasiliensis, F. merguiensis, Penaeus
Survival of shrimp varied from 48% (15‰) to 65% (35‰), showing semisulcatus and Fenneropenaeus indicus (Table 3). Nevertheless, the
higher values in treatments 25 and 35‰, but not differing significantly optimal salinity for growth and survival was lower for L. vannamei, F.
from the salinity 5‰ (Table 1). The food consumption analysis paulensis and Farfantepenaeus aztecus (Table 3). Although penaeids
indicated a significantly lower value in treatment 5‰, but not are resistant to salinity decreases during early life stages, most
significantly different from 15‰. Shrimp reared in the salinities of juveniles present higher growth rates in salinities between 25 and
25 and 35‰ showed the highest food consumption (Table 1). 35‰, which corresponds to migrations from estuarine areas back to
the sea that usually occur during this stage (Sang and Fotedar, 2004).
3.3. Haemolymph osmolality In addition to the physiological stress, growth of penaeids can be
affected in low salinities by the reduction in the assimilation of
The mean values of haemolymph osmolality in treatments from 15 nutrient (Bray et al., 1994). This fact was observed by Diaz (1995) for
to 25‰ were significantly different among them, but differed F. paulensis juveniles that required higher levels of protein in low
significantly from treatments 5, 10, 30 and 45‰ (Table 2). Haemo- salinity (10‰) when compared to those reared in salinities above
lymph osmolality ranged from 238 to 752.50 mosM kg− 1, which 20‰. According to Chen et al. (1992), F. chinensis reared in extreme
corresponded to salinities of 5 and 45‰, respectively. The medium salinities (5–10‰ and 30–40‰) need more time to complete their
osmolality differed significantly between all treatments. molting process, resulting in a reduction of feeding activity. Similar
The regression analysis between haemolymph and medium results were found by Chien (1992) and Staples and Heales (1991) for
osmolality (R2 = 0.9540) used to estimate the isosmotic point can P. monodon and F. merguiensis, respectively. Although the present
be summarized by the equation: study did not analyze the molting period, it may have influenced the
lower food consumption of F. subtilis in salinity 5‰.
Y = 236:0485 + 0:374X The osmoregulatory process is an important mechanism of
adaptation used by most crustaceans to overcome changes in salinity,
where Y = haemolymph osmolality (mosM kg− 1) and X = medium especially in estuarine and coastal marine environments (Pequeux,
osmolality (mosM kg− 1). 1995). Osmoregulatory capacity can be used as a tool to evaluate the
The isosmotic point was equivalent to 377.07 mosM kg− 1, which physiological conditions of cultured shrimp or their capacity for
corresponds to salinity 14‰. resistance to salinity stress (Charmantier et al., 1989). In this study, F.
subtilis hypoosmoregulate when salinity was above its isosmotic point
4. Discussion and hyperosmoregulate when salinity was below this point. This
adaptation is typical of penaeid shrimps and of most crustaceans that
Postlarvae of F. subtilis inhabit estuarine areas, and are thus inhabit estuarine areas (Mantel and Farmer, 1983; Lemaire et al.,
adapted to an environment in which salinity levels vary constantly. 2002).
Accordingly, our findings indicate that F. subtilis (PL50) presents a

Table 3
Table 2 Optimum salinity for growth and survival of some species of penaeids.
Mean (± SE) values of haemolymph and water osmolality (mosM kg− 1) of the shrimp
F. subtilis maintained in salinities ranging from 5 to 45‰ during 24 h. Species Development Salinity Reference
stages (‰)
Salinity (‰) 5 10 15 20 25 30 45
F. chinensis Juveniles 20–30 Chen et al. (1996)
Shrimp (n = 5) F. californiensis Postlarvae 33 Martinez et al. (1996)
Mean 238.0a 333.0b 442.6c 466.0c 476.0c 541.5d 752.5e F. brasiliensis Juveniles 35 Brito et al. (2000)
S.E. 8.0 6.0 17.7 9.0 14.5 7.5 17.5 F. merguiensis Juveniles 25 Ruttanagosrigit and Musig, (1982)
P. latisulcatus Juveniles 22–34 Sang and Fotedar, (2004)
Water (n = 3) P. indicus Postlarvae 17–35 Vijayyakumaran (1999)
Mean 138.7a 290.3b 398.0c 531.3d 672.0e 822.3f 1418.6g L. vannamei Juveniles 5–15 Bray et al. (1994)
S.E. 0.3 0.8 0.6 0.3 0.6 1.3 0.3 F. paulensis Postlarvae 10 Tsuzuki et al. (2003)
F. aztecus Juveniles 8.5–17 Venkataramiah et al. (1973)
Different superscript letters in the same row indicate significant differences (p b 0.05).
 E. Silva et al. / Aquaculture 306 (2010) 352–356
Chen and Lin (1998) and Brito et al. (2000) observed a close
relationship between haemolymph osmolality and medium osmolal-
ity in Litopenaeus stylirostris and F. brasiliensis, respectively. This study
confirms this relationship, where the haemolymph osmolality
increased with increasing medium osmolality. The slope obtained
by the linear regression between haemolymph and medium osmo-
lality was used to determine the osmoregulatory capacity of
F. chinensis (Chen et al., 1995; Chen and Lin, 1998), P. monodon
(Ferraris et al., 1986b) and P. latisulcatus (Sang and Fotedar, 2004).
The closer the slope to the isosmotic line (slope = 1), the lower the
capacity of shrimp to regulation (Chen et al., 1995). Based on the slope
value found in this study (0.374), F. subtilis can be considered a more
efficient regulator than F. chinensis (Chen and Lin, 1998), Farfantepe-
naeus duorarum and F. aztecus (Castille and Lawrence, 1981), but less
efficient than Litopenaeus setiferus (Rosas et al., 1999a), F. indicus
(Parado-Estepa et al., 1987) and L. vannamei (Castille and Lawrence,
1981). Furthermore, penaeids with very low slope values (e.g.
L. vannamei) have been reared in low salinity conditions (Laramore
et al. 2001; McGraw et al. 2002; Atwood et al. 2003; Saoud et al. 2003).
The isosmotic point to F. subtilis was estimated at 377.07 mosM kg− 1,
which is equivalent to salinity 14‰. The salinity of the isosmotic point has
been associated with the better condition for growth of penaeid shrimps
due to the supposed physiological comfort and lower energy expenditure
with osmoregulatory activity. Nevertheless, some species present a
superior growth performance in salinities above their isosmotic point
such as F. subtilis (this study), F. brasiliensis (Brito et al., 2000) and Penaeus
esculentus (O'Brien, 1994), while others improve growth close to their
isosmotic point such, as F. indicus, P. monodon and P. semisulcatus (Raj and
Raj, 1982), or even below this point, as observed for F. aztecus
(Venkataramiah et al., 1974), L. setiferus (Rosas et al., 1999b) and
L. vannamei (Bray et al., 1994).
The influence of salinity on the growth of penaeids has been
associated to combined factors of ecology, where the salinity influence
on the quantity and quality of natural food; physiology, through
osmoregulation processes; and even nutrition (Bray et al., 1994).
These factors were probably responsible for the lower growth of
F. subtilis found in this work in salinity 5‰. Moreover, under our
experimental conditions the growth performance of F. subtilis was
improved above its isosmotic point at the salinities of 25 and 35‰.
Acknowledgements
We thank Leônidas Oliveira for his assistance in running the trials
and Dr. Ronaldo Cavalli for his review of the manuscript. This work
was supported by the Pernambuco State Council of Science and
Technology (FACEPE) and the Brazilian Council for Scientific and
Technological Development (CNPq). S. Peixoto is research fellow of
CNPq.
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