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Nursery of Litopenaeus vannamei Reared in a Biofloc System: The Effect of

Stocking Densities and Compensatory Growth


Author(s): Wilson Wasielesky , Jr., Charles Froes , Geraldo Fóes , Dariano Krummenauer , Gabriele Lara
and Luis Poersch
Source: Journal of Shellfish Research, 32(3):799-806. 2013.
Published By: National Shellfisheries Association
DOI: http://dx.doi.org/10.2983/035.032.0323
URL: http://www.bioone.org/doi/full/10.2983/035.032.0323

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Journal of Shellfish Research, Vol. 32, No. 3, 799–806, 2013.

NURSERY OF LITOPENAEUS VANNAMEI REARED IN A BIOFLOC SYSTEM: THE EFFECT


OF STOCKING DENSITIES AND COMPENSATORY GROWTH

WILSON WASIELESKY, JR.,* CHARLES FROES, GERALDO FÓES, DARIANO KRUMMENAUER,


GABRIELE LARA AND LUIS POERSCH
Universidade Federal do Rio Grande (FURG), Instituto de Oceanografia, Laboratório de
Carcinocultura. Rua do Hotel, 02, Praia do Cassino, Rio Grande, Estado do Rio Grande do Sul,
Brasil, CEP 96210-030

ABSTRACT The purpose of this study was to evaluate the effect of stocking density and the occurrence of compensatory
growth in the white shrimp Litopenaeus vannamei in the nursery phase reared in a biofloc culture system. Two experiments were
performed in a recirculating water system with 12 experimental tanks (microcosm), each with a bottom area of 0.5 m2. The tanks
were supplied by a matrix 70-m2 biofloc technology water tank (macrocosm), where a shrimp biofloc technology systems culture
was maintained concurrently. The water was returned to the system by gravity, providing a daily renewal rate of approximately
4,800%. During the first experiment, L. vannamei postlarvae with an initial weight of 0.003 g were cultured for 30 days at densities
of 1,500, 3,000, 4,500, and 6,000 shrimp/m2. The cultures reached significantly different final weights (P < 0.05) of 0.45 g, 0.33 g,
0.30 g, and 0.23 g, respectively. Mean survival rates were greater than 87.6% in all treatments. To evaluate the occurrence of
compensatory growth in L. vannamei after a period of enclosure stress, the second experiment was organized in 2 phases. During
the first, shrimp postlarvae were reared at densities of 1,500, 3,000, 4,500, and 6,000 shrimp/m2 for 35 days. The shrimp were then
restocked at a density of 300 shrimp/m2 and were maintained until compensatory growth was observed. In 20 days, the shrimp
showed full compensatory growth. The mean final weight and survival rate did not differ significantly (P > 0.05) between the
treatments. The results confirm that the strategy of enclosure rearing can be applied to the culture of L. vannamei.

KEY WORDS: nursery, bioflocs, compensatory growth, Litopenaeus vannamei, stocking densities

INTRODUCTION to the growth and survival of the shrimp (Moss & Moss 2004,
Krummenauer et al. 2011). The choice of the most appropriate
Worldwide, marine shrimp culture has shown rapid growth
stocking density in the culture is critical to the economic
during the past few decades. Marine shrimp remains one of
viability of farms, exerting a considerable influence on system
the biggest commodities in aquaculture, and the Pacific white
productivity (Jackson & Wang 1998). Moreover, superintensive
shrimp Litopenaeus vannamei is the most widely reared species
of crustacean (FAO 2012). However, the need to maintain high culture involves the use of high stocking densities in small
productivity and reduce environmental impacts, in addition to culture units (Wasielesky et al. 2006, Krummenauer et al. 2011)
biosecurity considerations, has motivated studies of new culture and has a direct influence on the increasing production of
methods that may be applied with economic and environmental microbial flocs (Samocha et al. 2007).
sustainability (Sandifer & Hopkins 1996, Boyd & Clay 2002, Studies of different species of penaeid shrimp confirm that
Burford et al. 2003). the use of nurseries in biofloc systems contributes to the rapid
In this context, an interest in superintensive shrimp culture growth of the cultured organisms (Arnold et al. 2009, Fóes et al.
with minimal or no water exchange in biofloc technology (BFT) 2011, Emerenciano et al. 2012, Viau et al. 2012). In nursery
systems has emerged. In BFT system, highly oxygenated ponds phase, higher stocking densities are used and production costs
are fertilized with carbon-rich sources to stimulate the appear- are reduced. This system increases the number of crops in
ance of a predominantly heterotrophic bacterial biota (Ebeling temperate areas. In addition, it provides greater disease control
et al. 2006, De Schryver & Verstraete 2009). The bacteria that and increased survival at the end of the growth period (Kumlu
inhabit bioflocs assimilate the dissolved nitrogen compounds in et al. 2001, Yta et al. 2004, Fóes et al. 2011). However, the
the water, which are generated primarily by shrimp excretion excessive increase in stocking density may decrease the growth
and the decomposition of organic matter (Crab et al. 2007, and survival of the shrimp. This effect is associated with
De Schryver et al. 2008). This aquaculture practice enables the a combination of factors, including decreased viable space
recycling of the culture water through several cycles, making the and availability of natural food (Peterson & Griffith 1999),
system environmentally friendly (Avnimelech 1999, Krummenauer increased cannibalism (Abdussamad & Thampy 1994), degra-
et al. 2012). Furthermore, the stimulation provided by the biofloc dation of water quality (Nga et al. 2005), and the accumulation
can be an important feed supplement in the shrimp diet, contrib- of organic matter in the bottom of the tank (Arnold et al. 2006).
uting to digestion and protein retention, and the scope of these Nevertheless, many authors have reported that organisms
benefits includes the nursery phase (Tacon et al. 2002, Cuzon may exhibit compensatory growth followed by increased
et al. 2004, Otoshi et al. 2011, Xu et al. 2012). growth if they are farmed under suitable conditions (Nicieza
Among the many aspects of the management of shrimp & Metcalfe 1997, Ali et al. 2003, Oh et al. 2007). This increase in
farming in biofloc systems, the stocking density is related directly growth rates is the result of strategies adopted by the organisms,
including a higher food intake in the poststress period (hyper-
*Corresponding author. E-mail: manow@mikrus.com.br phagia), increased feed efficiency, reduced metabolic wastes,
DOI: 10.2983/035.032.0323 and reduced locomotion (Quinton & Blake 1990, Jobling et al.

799
800 WASIELESKY ET AL.

1994, Ali et al. 2003). Wu et al. (2000), and Wu and Dong (2001, A Litopenaeus vannamei biofloc culture was maintained in
2002) report compensatory growth in Fenneropenaeus chinensis the macrocosm tank without water renewal. The mean weight
after periods of food restriction, a protein-deficient diet, and of the shrimp in the macrocosm tank ranged from 3 to 16 g
low temperature. throughout the study, and the stocking density used in this tank
However, no previous studies have correlated stocking was 300 shrimp/m2. Fertilization with molasses was performed
densities and compensatory growth in penaeid shrimp in to maintain a C:N ratio of 6:1 (Ebeling et al. 2006, Samocha
combination with the possible benefits of biofloc culture during et al. 2007, Avnimelech 2009). To maintain adequate pH levels
the nursery phase. Thus, the objective of this study was to eval- for shrimp culture, sodium carbonate (soda ash; pH+ from
uate the best stocking density for nursery culture of Litopenaeus HidroAll Ltda, São Paulo, Brazil) was applied as proposed by
vannamei in a BFT system and, subsequently, to evaluate the Furtado et al. (2011).
effect of different stocking densities on the compensatory
growth of shrimp under the same culture conditions. Experimental Design

MATERIALS AND METHODS Trial 1: Use of Different Stocking Densities During the nursery phase of
Litopenaeus vannamei in a Biofloc Culture System
Location The experimental tanks were stocked with Litopenaeus
vannamei postlarvae (PL 10) acquired from a commercial lab-
The experiments were performed at the Marine Station of
oratory (Aquatec Ltd., Canguaretama, Rio Grande do Norte,
Aquaculture, Institute of Oceanography, Federal University of
Brazil). The initial mean weight was 0.003 g (±0.001).
Rio Grande.
The experimental design was randomized with 4 treatments
corresponding to 4 stocking densities: 1,500 (T1500), 3,000
Experimental Units and Environmental Conditions
(T3000), 4,500 (T4500) and 6,000 (T6000) shrimp/m2, with
To evaluate the specific effects on shrimp performance of 3 replicates each. The shrimp were 4 four times a day (0900 HR,
stress caused by population density, a water biofloc recircula- 1200 HR, 1700 HR, and 2000 HR) with commercial feed (Potimar
tion system (Fig. 1) was designed in a greenhouse. This system 40J–40% CP). The feed was adjusted according to Jory et al.
was used because the main objective of the experiment was to (2001). The experiment lasted 30 days.
observe behavioral aspects of shrimp in terms of tolerance to
a crowding effect. Furthermore, this experimental design aimed Trial 2: The Effect of the Stocking Density During the Nursery Phase
Under Compensatory Growth of Litopenaeus vannamei in a Biofloc
to maintain the same water quality and the amount of natural
Culture System
food (bioflocs) to detect the maximum stocking density for the
nursery phase. The system included twelve 180-L tanks (micro- The second study was performed in 2 phases. During the
cosms) with a bottom area of 0.5 m2, and intense and individual first, postlarvae of Litopenaeus vannamei with an initial mean
aeration. All the tanks had water output driven by gravity into weight of 0.002 g (±0.001) were nursed for 35 days at stocking
a return pipe. The water was directed into a 100-m3 raceway densities of 1,500, 3,000, 4,500 and 6,000 shrimp/m2. After the
tank (macrocosm), where the air supply was introduced with first 35 days (nursery phase), the nursed shrimp from different
a blower. The water was pumped back into the experimental stocking densities were stocked in the same tanks but at a
tanks, with a flow rate of 6.6 L/min per tank (4,800% stocking density of 300 shrimp/m2 (growth phase), with 4 rep-
recirculation per day). Prior to initiating the experiment, a test licates each. The shrimp were fed twice a day (0900 HR and
was performed to determine whether it was possible to maintain 1700 HR) with commercial feed (Potimar Active 38–38% CP).
the water quality of the experimental tanks under the same The feed of each treatment was adjusted according to Jory et al.
conditions as the matrix tank. The results of this test showed (2001). Compensatory growth was assessed according to the
that the factor of water quality did not interfere with the shrimp patterns described by Jobling et al. (1994). The second phase
at different stocking densities. was performed until the first indication of full compensatory
growth in the shrimp reared.

Water-Quality Parameters

Temperature, pH, dissolved oxygen, and salinity were mea-


sured daily in each experimental tank with a multiparameter
analyzer (model 556 MPS; YSI Inc.). Total ammonia was
measured according to UNESCO (1983). Nitrite, nitrate, and
phosphate were measured according to Strickland and Parsons
(1972). These analyses were conducted every 3 days and only
in the matrix tank.
Water samples from the matrix tank were collected weekly to
measure chlorophyll a. To extract the photosynthetic pigment,
90% acetone (Merck, PA) was used in the dark at –12!C for
24 h. The chlorophyll concentrations were then determined by
fluorometry (Welschmeyer 1994). The total suspended solids
(measured in milligrams per liter) were also analyzed weekly
Figure 1. Diagram of biofloc recirculation system. (adapted from Strickland and Parsons (1972)). Solids that could
NURSERY AND COMPENSATORY GROWTH IN A BFT SYSTEM 801

settle (measured in milliliters per liter) were quantified using TABLE 2.


Imhoff cones, in which samples from 1 L of water were deposited Mean values (%SD), with minima and maxima, of the
for 15 min. The settled volume was then read (Avnimelech 2007). water-quality parameters from the matrix tank recorded
during the experimental period.
Experimental Management and Shrimp Performance
Parameter Mean % SD Minimum Maximum
In both experiments, initial biometric measurements were
made on 200 randomly selected shrimp for initial mean weight Salinity 34.3 ± 0.34 33 36
Transparency (cm) 13.7 ± 2.2 9.5 16.2
determination. After the beginning of the study, 50 animals
Total suspended solids (mg/L) 273 ± 128.82 123 437
from each tank were weighed every 10 days in experiment 1 and
Solids that could settle (mL/L) 24.7 ± 6.3 17.42 32.3
weekly in experiment 2. The shrimp were returned to their Chlorophyll a (mg/L) 204.3 ± 31.2 136.1 187.6
original tanks after weighing. Total ammonia (N-AT, mg/L) 0.19 ± 0.06 0.04 0.53
The weight gain was calculated using the following formula: Nitrite (N-NO2, mg/L) 2.85 ± 1.9 1.45 6.79
Weight gain ¼ Final average weight – Initial weight. The daily Nitrate (N-NO3, mg/L) 7.65 ± 2.54 3.19 11.37
specific growth rate (G%) was calculated according to Bagenal Phosphate (P-PO4, mg/L) 0.17 ± 0.05 0.09 1.07
(1978) from the following formula: G% ¼ ðWf # WiÞ 3 100=ND,
where Wf is the final weight, Wi is the initial weight, and ND
is the number of days of culture. The feed conversion rate
(FCR) was calculated from the following formula: FCR ¼ 0.45 ± 0.12 g, 0.33 ± 0.09 g, 0.3 ± 0.06 g, and 0.23 ± 0.09 g,
Foodoffered=Biomassincrement: respectively (Fig. 2, Table 3).
Survival was calculated as Survivalð%Þ ¼ ðTotalweight= The means of the final weight and daily specific growth rate
IndividualweightÞ 3 100: The data were transformed (arcsine of in T3000 and T4500 did not differ significantly. However,
the square root) before analysis. T1500 and T6000 differed, showing the highest and lowest
After the homoscedasticity and normality of the data were average, respectively. For T1500, T3000, and T4500, the FCR
verified, the data were analyzed with 1-way analysis of variance did not show statistically significant differences among the
and with a Tukey test. The differences were considered signif- treatments, ranging from 0.95 ± 0.04 to 1.11 ± 0.05. The FCR
icant at 95%. The results are presented as the mean ± SD. was higher in T6000 (1.61 ± 0.03) than in the other treatments.
The highest productivities were 1.23 kg/m2 in T4500 and 1.20
kg/m2 in T6000. Treatments T1500 and T3000 produced final
RESULTS biomass values of 0.65 kg/m2 and 0.94 kg/m2, respectively.
These values differed statistically from those found for the other
Trial 1
treatments (Table 3).
The temperature, salinity, pH, and dissolved oxygen did not
differ significantly among the treatments (Table 1). Table 2 Trial 2
shows the mean values (with minima and maxima) of the water- Temperature, salinity, pH, and dissolved oxygen did not
quality parameters observed in the matrix tank, in which a water differ significantly among the treatments (Table 4). Table 5
biofloc recirculation system was operated. shows the mean values (with minima and maxima) of the water-
A higher percentage of survival was observed in T1500 (96.26 ± quality parameters observed in the matrix tank, in which a water
2.26%). No significant difference was observed between treat- biofloc recirculation system was operated.
ments T3000 and T4500 (with means of 95.55 ± 6.13% and 95.49 ± During the first phase of trial 2, after 35 days of the nursery
2.23%, respectively). The lowest survival was observed in T6000 phase, T1500 showed a greater final weight (0.91 ± 0.27 g), with
(87.6 ± 5.16%), differing significantly (P < 0.05) from the other significant differences from the T3000 and T4500 treatments,
treatments (Table 3). which showed averages of 0.59 ± 0.35 g and 0.55 ± 0.31 g,
At the end of the 30 days of the study, the shrimp stocked in respectively (Fig. 3A). The lowest average weight was 0.38 ±
T1500, T3000, T4500, and T6000 showed mean final weights of 0.18 g and it occurred in T6000. This value differed from those
found in the other treatments. At the end of the nursery phase,
TABLE 1. shrimp originating from different densities and restocked to a
density of 300 shrimp/m2 (i.e., T1500 (n ¼ 300), T3000 (n ¼ 300),
Mean values (%SD) of water-quality parameters recorded in T4500 (n ¼ 300), and T6000 (n ¼ 300)) showed no difference (P >
Litopenaeus vannamei nursed in a biofloc system at different
0.05) among the treatments in any of the zootechnical parameters
stocking densities (1,500, 3,000, 4,500, and 6,000 shrimp/m2).
after the 20th day of experimental fattening. This result indicates
that the shrimp reached full compensatory growth (Fig. 3B,
Treatments (trial 1) Table 6).
Parameter T1500 T3000 T4500 T6000
DISCUSSION
Temperature 29.05 ± 0.82 29.5 ± 0.95 29.05 ± 0.84 29.45 ± 0.95
(!C) The mean values of the water-quality parameters recorded
pH 7.56 ± 0.33 7.43 ± 0.41 7.61 ± 0.45 7.59 ± 0.39 during the 2 trials showed no effect on the growth and survival
Dissolved 6.28 ± 1.31 6.58 ± 1.57 6.11 ± 1.23 5.97 ± 1.07
of the shrimp. The microcosm system maintained the water-
oxygen
(mg/L)
quality parameters at the same basic conditions in all treatments
as a result of the high recirculation rate. The experimental
802 WASIELESKY ET AL.

TABLE 3.
Mean (%SD) of final weight, survival, feed conversion rate (FCR), specific daily growth rate (G%), final biomass, and final density
of postlarvae of Litopenaeus vannamei nursed at densities of 1,500, 3,000, 4,500, and 6,000 shrimp/m2 in a biofloc system.

Treatments (trial 1)
Parameter T1500 T3000 T4500 T6000
Initial weight (g) 0.003 ± 0.001 0.003 ± 0.001 0.003 ± 0.001 0.003 ± 0.001
Final weight (g) 0.45 ± 0.12a 0.33 ± 0.13b 0.30 ± 0.09b 0.23 ± 0.09c
Survival (%) 96.26 ± 2.26a 95.55 ± 6.13a 95.49 ± 2.23a 87.6 ± 5.16b
FCR 0.95 ± 0.04a 1.03 ± 0.07a 1.11 ± 0.05a 1.61 ± 0.03b
G (%) 1.49 ± 0.19a 1.09 ± 0.16b 0.99 ± 0.17b 0.76 ± 0.10b
Productivity (kg/m2) 0.65 ± 0.03a 0.94 ± 0.04b 1.23 ± 0.07c 1.20 ± 0.09c
Final density (shrimp/ m2) 1,444 ± 28.07a 2,886 ± 173.9b 4,297 ± 85.16c 5,256 ± 262.78d

Different superscript letters within the same column indicate significant differences (P < 0.05).

design, based on that proposed by Moss and Moss (2004) and et al. 2010). The total suspended solids and solids that could
Wasielesky et al. (2006), showed efficiency in separating the settle were maintained at levels recommended by Samocha et al.
effects of population stress and the degradation of water quality (2007) and Gaona et al. (2011).
on the performance of the shrimp. Stocking density is a factor that affects the growth and
The dissolved oxygen was maintained above recommended productivity resulting at the end of the culture of many species,
levels, and the minimum concentration observed was 3.8 mg/L. and an inverse relationship between these factors is generally
During the 2 experiments, the temperature was maintained observed (Sturmer & Lawrence 1987, Moss & Moss 2004, Fóes
within the optimum range for the rearing of the species (Wyban et al. 2011, Krummenauer et al. 2011). However, many studies
et al. 1995). Maicá et al. (2012) recommended salinity values report that survival is not affected by the stocking density in
above 25 ppm for Litopenaeus vannamei culture in a BFT sys- most cases, primarily if biofloc culture is used. Our results
tem. In the current study, this parameter was maintained within confirm the substantial support capacity of the system at high
acceptable levels for this culture. The use of sodium carbonate, stocking densities (Cohen et al. 2005, Mishra et al. 2008, Fóes
as proposed by Furtado et al. (2011), helped to keep the pH et al. 2011).
values suitable for good growth performance of the species In the first study, it was observed that the mean final weight
(Van Wyk & Scarpa 1999). and survival in T6000 were significantly less than those observed
The total ammonia, nitrite, and nitrate in the matrix tank in the other treatments, even though the final productivity was
remained below toxic levels for Litopenaeus vannamei (Van high. Arnold et al. (2006) identified survival as the primary pa-
Wyk & Scarpa 1999, Lin & Chen 2001, Lin & Chen 2003). The rameter to be considered in nursery cultures of Penaeus monodon.
total ammonia was maintained at low levels during the exper- In the light of these findings on the relationship between
iments, most likely resulting from the development of the stocking density and survival in the cultured shrimp, a density
microbial community that was established in the culture water. of 6,000 shrimp/m2 is unsuitable for commercial production,
Because of the addition of molasses and the C:N ratio adjust- given the substantial investment needed to acquire the post-
ment, the bacterial community was able to use the dissolved larvae and the feed during the growing season. The high levels
nitrogen to form biomass (Bratvold & Browdy 2001, Ballester of survival in treatments T1500, T3000, and T4500 were related
to the high water quality that was maintained during the study
and also to the availability of in situ production of additional
feed provided by the bioflocs (Crab et al. 2012).

TABLE 4.
Mean values (%SD) of water-quality parameters recorded in
Litopenaeus vannamei nursed in a biofloc system at different
stocking densities (1,500, 3,000, 4,500, and 6,000 shrimp/m2)
and after restocking at 300 shrimp/m2.

Treatments (trial 2)
Parameter T1500 (300) T3000 (300) T4500 (300) T6000 (300)
Temperature 28.15 ± 1.06 28.45 ± 1.18 27.85 ± 0.86 28.35 ± 1.15
(!C)
pH 7.52 ± 0.34 7.63 ± 0.36 7.51 ± 0.48 7.64 ± 0.54
Dissolved 5.38 ± 1.31 5.17 ± 1.62 5.24 ± 1.51 5.11 ± 1.81
Figure 2. Growth of Litopenaeus vannamei nursed in different stocking oxygen
densities (1,500, 3,000, 4,500, and 6,000 shrimp/m2) for 30 days in (mg/L)
a biofloc culture system (trial 1).
NURSERY AND COMPENSATORY GROWTH IN A BFT SYSTEM 803

TABLE 5. did not differ among these treatments. T6000 showed a greater
Mean values (%SD), with minima and maxima, of the water- FCR, which indicates that this parameter is influenced by the
quality parameters for the matrix tank recorded during the high stocking densities, and it also shows that the ability of
experimental period. shrimp to graze on the microbial community may decrease or be
less efficient at these high densities.
A decrease in the growth and survival of juvenile penaeids at
Parameter Mean % SD Minimum Maximum
high stocking densities is related to a combination of factors,
Salinity 35.3 ± 0.72 32 36 such as the decrease in the availability of space and natural
Transparency (cm) 11.7 ± 3.3 8.5 17.1 productivity, in addition to degradation of water quality and
Total suspended solids (mg/L) 302 ± 164.19 122 444
accumulation of anaerobic sediment (Maguire & Leedow 1983,
Solids that could settle (mL/L) 28.6 ± 6.3 16.9 35.5
Peterson & Griffith 1999, Moss & Moss 2004). In the current
Chlorophyll a (mg/L) 157.3 ± 22.32 136.1 187.6
Total ammonia (N-AT, mg/L) 0.34 ± 0.06 0.11 0.89 study, the water quality and the availability of natural food
Nitrite (N-NO2 , mg/L) 2.72 ± 1.7 1.21 7.93 were similar for all treatments because the biofloc recirculation
Nitrate (N-NO3, mg/L) 9.89 ± 2.54 6.93 17.66 system used was the same for all tanks. Thus, the specific growth
Phosphate (P-PO4 mg/L) 0.23 ± 0.1 0.04 1.23 rate was influenced by the space limitation to which the shrimp
were subjected, showing an inverse relationship with an increase
in stocking density. According to the results obtained in trial 1,
it is possible to nurse Litopenaeus vannamei in a BFT system at
The natural food source provided by the growth of the
stocking densities of up to 4,500 shrimp/m2 with no reduction in
microbial community offers a high-quality food supplement
the survival of the cultured shrimp. Nevertheless, the high
that benefits the shrimp, especially during the nursery phase
stocking densities have a negative influence on the final weight
(Emerenciano et al. 2012, Godoy et al. 2011). Because of the
of the shrimp.
improved utilization of natural productivity, a decreased FCR
The compensatory growth in fish and crustaceans subjected
was expected (Wasielesky et al. 2006, Mishra et al. 2008), and
to stress usually occurs at intervals of 10–30 days. This com-
this effect is also reflected by the specific growth rate of the
pensatory growth response may be classified into the following
shrimp. The FCRs were lower in T1500, T3000, and T4500, and
3 categories: (1) partial compensation, in which the animals
cannot reach the control group level at the same age as un-
restricted contemporaries; (2) full compensation, in which the
animals reach the weight of the control group at the same age as
the control group; and (3) overcompensation, in which the
animals are larger than the control group during the same time
frame (Jobling et al. 1994, Ali et al. 2003). During the second
experiment, following the same pattern of growth that occurred
during the first study, the shrimp that had spent 35 days in the
nursery at higher densities were restocked at 300 shrimp/m2
with an initial weight less than the shrimp in the T1500 (n ¼ 300)
treatment. At the end of 20 days, during which animals from
all treatments were reared at 300 shrimp/m2, there were no
significant differences in any parameter of growth performance.
These results indicate that full compensatory growth had
occurred.
Although the daily specific growth rate did not show
statistical differences among treatments, these values were
sufficient to recuperate the growth deficit caused by the high
stocking densities used during the nursery phase. Wu et al.
(2001) considered the daily specific growth rate an adequate
parameter for the detection of compensatory growth. Several
authors affirm that compensatory growth is triggered by
hyperphagia in response to a period of food shortage, increasing
the appetite level (Russell & Wootton 1992, Nicieza & Metcalfe
1997). Despite that fact that the FCRs were equal for different
treatments (P > 0.05), the amount of supplied feed for each
treatment was different (P < 0.05). This occurred because
shrimp were fed according tables proposed by Jory et al.
(2001), which recommend different feeding rates for each
shrimp size. Furthermore, because the BFT system may in-
Figure 3. (A, B) Nursery of Litopenaeus vannamei in different stocking crease the growth performance and protein utilization of the
densities (1,500, 3,000, 4,500, and 6,000 shrimp/m2) for 35 days in shrimp (Ballester et al. 2010), we should not disregard the
a biofloc culture system (A) and then restocked in the same stocking importance of natural productivity in the improved utilization
density (300 shrimp/m2) to verify the compensatory growth for more than of the food, even after a period of great stress caused by high
20 days (B) (trial 2). population densities.
804 WASIELESKY ET AL.

TABLE 6.
Mean (%SD) of the final weight, survival, feed conversion rate (FCR), daily specific growth rate (G%), final biomass, and final
density of Litopenaeus vannamei nursed in a biofloc system at different stocking densities (1,500, 3,000, 4,500, and 6,000 shrimp/m2)
for 35 days and after restocking at 300 shrimp/m2 for 20 days, when they achieved full compensatory growth.

Treatments (trial 2)
Parameters T1500 (300) T3000 (300) T4500 (300) T6000 (300)
a b b
Initial weight (g) 0.91 ± 0.27 0.59 ± 0.35 0.55 ± 0.31 0.38 ± 0.18c
Final weight (g) 3.70 ± 1.07 3.62 ± 1.02 3.83 ± 0.94 3.60 ± 1.21
Survival (%) 93.26 ± 4.66 96.55 ± 3.47 98.14 ± 4.11 93.49 ± 4.32
FCR 1.54 ± 0.11 1.24 ± 0.17 1.32 ± 0.22 1.28 ± 0.37
G (%) 12.95 ± 2.35 15.15 ± 3.21 16.4 ± 3.54 16.4 ± 3.54
Productivity (kg/m2) 1.04 ± 0.05 1.05 ± 0.03 1.13 ± 0.04 1.01 ± 0.04
Final density (shrimp/m2) 279.78 ± 11.1 289.65 ± 8.67 294.42 ± 11.3 280.47 ± 11.2

Different superscript letters in a row means significant differences.

Freetly et al. (1995) suggest that the compensatory growth units to enhance productivity through the maximum use of
response is a physiological mechanism that increases the available space and improvement of management techniques.
fraction of energy directed to growth. This process occurs by
increasing the efficiency of feed utilization and decreasing CONCLUSIONS
energy loss through feces and ecdysis (Dobson & Holmes
1984, Wei et al. 2008). This response appears to have occurred The results of this study confirm that increased stocking
in the current study. The high survival in all treatments at the density during the nursery phase affected the growth and
end of the second experiment is also a good indication that the survival of Litopenaeus vannamei postlarvae nursed in a biofloc
bioflocs contributed to the health of the shrimp even after system. If the aim of the farmer is to achieve rapid growth
exposure to high densities. Crab et al. (2012) state that bacteria during this phase, a use of 1,500 shrimp/m2 is suggested.
and their products may have immunostimulatory effects on However, if the area of the farm is restricted, nursery culture
animal growth. This effect can increase the survival and re- can be performed at stocking densities up to 4,500 shrimp/m2
sistance of animals, even during stressful situations. without losses in survival. Our results also indicate that the
The results of this study indicate that an increase in stocking biofloc system can contribute to achieving full compensa-
density reduced the growth rates and survival of shrimp during tory growth of shrimp within 20 days at a stocking density of
the nursery phase. These results agree with those presented by 300 shrimp/m2 during the growth phase.
Moss and Moss (2004), Krummenauer et al. (2011), and Otoshi
et al. (2007). However, the best productivity was obtained in the ACKNOWLEDGMENTS
treatment with a density of 4,500 shrimp/m2. These shrimp
showed compensatory growth after transfer to a lower density. We are grateful for the financial support provided by the
This result showed that a density of 4500 shrimp/m2 was slightly National Council for Scientific and Technological Develop-
better for the nursery of Litopenaeus vannamei in the BFT system. ment (CNPq), Ministry of Fishery and Aquaculture (MPA),
Farms for Litopenaeus vannamei using the BFT system and Coordination for the Improvement of Higher Level
involve high investments in cost and operation (Browdy et al. Personnel (CAPES). We especially thank Centro Oeste Racxões
2001, Boyd & Clay 2002, Wasielesky et al. 2006). The results of S.A. (GUABI) for donating the experimental diets. W. J.
this study may help in the design and planning of the culture Wasielesky and L. H. Poersch are research fellows of CNPq.

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