Ecological Modelling 437 (2020) 109291

Contents lists available at ScienceDirect

Ecological Modelling
journal homepage: www.elsevier.com/locate/ecolmodel

Empirical modeling of feed conversion in Pacific white shrimp (Litopenaeus T

vannamei) growth
Marcos Estevão Santiago de Melo Filhoa,b, Marco Shizuo Owatarib, , José Luiz Pedreira Mouriñoa,
⁎

Bruno Augusto Mattar Carciofia, Hugo Moreira Soaresa

a
Federal University of Santa Catarina Chemical and Food Engineering Department, Florianopolis, SC, Brazil
b
Federal University of Santa Catarina, Aquaculture Department, Florianopolis, SC, Brazil

ARTICLE INFO ABSTRACT

Keywords: Litopenaeus vannamei or Pacific white shrimp is one of the main shrimp species cultivated around the world.
Stoichiometric equation Despite its easy handling, there is no detailed description of its metabolic performance. This work developed a
Specific growth rate novel mathematical model for L. vannamei growth based on mass conservation and chemical equations for feed
Oxygen consumption oxidation, allowing to calculate shrimp specific growth rate, specific rate of feed consumption, specific rate of
Ammonia excretion
ammonia nitrogen excretion, and oxygen specific consumption rate. Results, when compared to literature data,
Statistical model
indicate a correlation higher than 99%, demonstrating that the proposed approach can successfully predict the
growth based on the assessed parameters. Through this strategy it is possible to optimize the management of
shrimp cultivation, especially in relation to dissolved oxygen, availability of feed and ammonia excretion, ac-
cording to each stage of shrimp growth. Based on the environmental conditions maintained during the present
study, the result of the present mathematical model can be a potential tool for reducing costs and environmental
impacts in aquaculture shrimp farm.

1. Introduction environment, affecting the metabolic rate (Regnault, 1987; Schmidt-

Marine shrimp are highly cultivated and commercialized, re-
presenting the second largest group of aquaculture species and are an
important economic source for many developing countries in Asia and
Latin America. This industry generates around 1.51 billion dollars US /
year (FAO, 2015, 2018). One of the main shrimp species cultivated is
the L. vannamei, well-known as Pacific white shrimp. Its high re-
productive potential, rapid growth, high market value, and the ex-
istence of an established methods package for its cultivation are some of
the characteristics that enabled the successful establishment of this
industry (FAO, 2014). Despite ease of handling, regulation of dissolved
oxygen, salinity, temperature and feed quality are crucial during the L.
vannamei cultivation (Ponce-Palafox et al., 1997a; Rosas et al., 2001;
Hernández et al., 2006; Valenzuela-Quiñónez et al., 2011).
Dissolved oxygen is the main limiting factor in aquaculture systems
(Vinatea, 1997) and during shrimp cultivation it is essential to maintain
adequate levels of oxygen, ranging from 4.0 to 5.5 mg of O2 L − 1
(Timmons and Ebeling, 2007). Changes in oxygen consumption reflect
physiological responses of the shrimp to environmental conditions,
showing the relationship of dependence between the animals and the
Nielsen, 2002). Inadequate conditions are potential stressors that ne-
gatively affect growth, increase susceptibility to diseases, causing
mortality (Bett and Vinatea, 2009). Abiotic factors such as water tem-
perature and salinity can directly influence animal physiology. Tem-
perature is related to the feed conversion rate and the Pacific white
shrimp reaches optimum growth in the range between 25 and 28 °C
(Ponce-Palafox et al., 1997a; Hernández et al., 2006). On the other
hand, salinity is related to osmoregulation process. Due to the ability of
L. vannamei to maintain osmotic regulation over a wide range, this
species can inhabit water with salt concentration values ranging from
0.5 to 40 ‰. However, the optimal range for cultivation is from 15 to
25 ‰ (Saoud et al., 2003; Jayasankar et al., 2009; Paz et al., 2011;
Maicá et al., 2014). According to Gao et al. (2016), under intensive
cultivation variations in salinity may break homeostasis and lead to
significant stress, slow growth, low survival rate, causing high eco-
nomic losses. The availability and quality of feed, measured by protein
value, directly affects shrimp production (Mente, 2003). Commercial
feeds contain from 30 to 50% of crude protein, composed mostly of
marine animal protein products such as fish, shrimp, and squid meal
(Mente et al., 2002). It is worth noting that feed composition directly

⁎
Corresponding author.
E-mail address: marco.owatari@posgrad.ufsc.br (M.S. Owatari).

https://doi.org/10.1016/j.ecolmodel.2020.109291
Received 22 March 2020; Received in revised form 10 September 2020; Accepted 13 September 2020
Available online 28 September 2020
0304-3800/ © 2020 Elsevier B.V. All rights reserved.
M.E.S. de Melo Filho, et al.
influences the nitrogen levels in the culture system (Lee and
Lawrence, 1985; Seidman and Lawrence, 2009).
The most common methodology used to maintain nitrogen balance
during shrimp cultivation considers the difference between the nitrogen
contained in the feed and that excreted into the water and within the
feces. The remaining nitrogen is that incorporated by shrimp. However,
the wide variation in nitrogen incorporation, 23% to 63%, make this
method practically ineffective (Boyd and Teichert-Coddington, 1995;
Hopkins et al., 1995; Velasco et al., 1999). An alternate method for
quantifying nitrogen incorporation is through the elementary mass
balance and this makes possible correlation, using stoichiometric bal-
ance, between the oxidation of the nitrogen within the feed and the
products of shrimp metabolism.
The modeling that can quantitatively represent an aquaculture
system and the way in which the interactions between the variables that
act in these systems occur, allow us to better understand and support
the production processes in their real context. For example, an equation
can correspond perfectly to describe the change in any biological
parameter associated with body weight due to allometric dependence
(Zotin, 2015). Representations that aim to fully understand ecosystem
phenomena offer an alternative to characterize biotic communities and,
initially, tend to be highly abstract. Several holistic hypotheses can be
tested in ecosystems. Macroscopic concepts can promote a better un-
derstanding of physical-biological interactions and suggest the devel-
opment of new instrumentation and technological methods
(Ulanowicz and Platt, 1985)
Mathematical models can and should be used to interpret the hy-
potheses from a quantitative point of view, giving us the information
needed to predict consequences of rearing conditions and highlight
conditions that should be corrected or maintained to obtain a produc-
tive shrimp growth cycle (Klemetson and Rogers, 1985; Chen and
Malone, 1992; Panchang et al., 1997; Sodré, 2007; Byron et al., 2011;
Timmons et al., 2001; Seginer, 2016). However, different mathematical
models can demonstrate better performance than others, providing
useful information for prediction of animal growth (Teleken et al.,
2017)
In this way, there is a need to better establish description and pre-
diction of metabolic performance with respect to feed consumption and
product generation in shrimp aquaculture systems. Thus, this work
developed an elemental balance model based on stoichiometric che-
mical equation of the feed oxidation, allowing the user to determine
specific growth and oxygen specific consumption rates for marine
shrimp L. vannamei.
2. Material and methods
The shrimp used in this study were taken from the stocks routinely
cultivated by the Shrimp Marine Laboratory (LCM) of the Federal
University of Santa Catarina (UFSC) for the purpose of maintaining
breeding stocks.
2.1. Sample preparation and storage
At the time of collection, rearing conditions were temperature 28.65
°C, salinity 34.5 psu (practical salinity unit), dissolved oxygen 6.18 mg
L − 1, pH 7.87, total ammonia 0.30 mg L − 1, Nitrate 1.26 mg L − 1 and
Alkalinity 143.31 mg CaCO3 L − 1.
The commercial feed used to grow shrimp (GUABI brand) had a
stated composition: 34.67 ± 0.5% crude protein, 7.5 ± 1.2% ethereal
extract, 0.5 ± 0.4% crude fiber, 13 ± 1.5% of mineral matter,
3 ± 0.2% calcium, 1.45 ± 0.3% of phosphorus, 95.48 ± 3.4% of dry
matter and 3259.3 ± 55.9 of energy (kcal). For elemental analysis a
sample of 60 gs of feed pellets were ground to obtain a homogeneous
flour and dried at 55 °C until constant weight.
Samples containing thirty shrimp were obtained from five cultiva-
tion tanks with populations at different stages of cultivation. The
2
Ecological Modelling 437 (2020) 109291
individual weight of the shrimp used was intentionally chosen between
2 and 10 g, in order to represent the different growth phases, making up
five categories, with weights of 1 to 2, 2 to 4, 4 to 6, 6 to 8 and 8 to 10 g.
From each weight category, two grams of muscle were removed from
the shrimps and a pool containing muscle from all weight categories
was created. In this way a unique elemental sample was generated
containing information with characteristics of all shrimp sizes. Samples
were minced and dehydrated through evaporation within a drying oven
at 55 °C until sample weights stabilized. Samples were then ground to a
homogeneous flour and dried again to constant weights at 55 °C.
A feces sample of 10 g was collected from five different shrimp
cultivation tanks (50 m3) from LCM/UFSC and oven dried at 55 °C to
constant weight, ground to a homogeneous flour and dried again to
constant weight at 55 °C. After dried, all samples were stored in a de-
humidifier to avoid rehydration.
2.2. Elemental analyses
Elemental analyses of the samples were performed on a Leco
CHNOS TruSpec Micro equipment, following the ASTM D 5373 method
(ASTM, 2012). The elemental analysis involves determining the mass
percentage of carbon, hydrogen, nitrogen and oxygen in the dried
samples of feed, shrimp biomass and feces. In this methodology, the
samples are burned in an oven at a temperature of 1423 K. The com-
bustion products are subsequently transferred to a secondary oven with
a temperature of 1123 K for total oxidation of the compounds. In this
technique, the amounts of carbon and hydrogen in the form of CO2 and
H2O were determined using an infrared detector. The nitrogen content
was measured using a thermal conductivity detector. The oxygen was
measured in the same equipment, by a separate module that performs
the combustion of the compound in a pyrolysis oven at a temperature of
1573 K, which guarantees the complete release of oxygen in the form of
CO2. The results of the analysis were obtained as a percentage and the
mass calculation of each element was performed. Shrimp biomass used
in TruSpec = 2.0573 ± 0.04 mg, mass of feces used in
TruSpec = 2.1357 ± 0.05 mg and mass of feed used in
TruSpec = 2.0717 ± 0.03 mg.
2.3. Stoichiometric balance
Eq. (1) is proposed for describing the L. vannamei growth based on
feed consumption and the main products generated.
Ingested feed + O2 = Shrimp Biomass + Feces + Urea + NH3 + H2 O
+ CO2 (1)
From the elemental analysis, we obtained the minimum formula
elemental composition (C, H, N, and O composition in whole molar
count) of each component of Eq. (1) necessary to balance the elemental
composition on both sides of the equation (Eq. (2), below).
The elemental balance budget, schematically represented in Fig. 1,
was estimated considering the percentage of dry matter digestibility
61.53% as stated by Buglione-Neto et al. (2013) for the same shrimp
species and feed. From the digested nitrogen, 20% is converted to
biomass, and 80% is not assimilated and converted into shrimp tissue,
being eliminated in the excretion, in which, the excretion budget is
feces (20%), ammonia-nitrogen (72%) and urea (8%) (Ebeling et al.,
2006).
From the nitrogen balance between undigested feed, feces, shrimp
biomass, ammonia, and urea described in Fig. 1, we determined the
coefficients b, c, d, and e of Eq. (2). Additionally, we calculated the
molar balance for carbon, hydrogen, and oxygen, and, through a system
containing three equations and three variables, we determined the
coefficients a, f, and g of Eq. (2).
M.E.S. de Melo Filho, et al. Ecological Modelling 437 (2020) 109291

Fig. 1. Nitrogen mass balance during shrimp culti-

vation according to Ebeling et al. (2006) based on
100 g of dry feed: 8.10 g of N are ingested, 4.984 g
are digested and assimilated, 0.99 g are converted to
biomass, 0.319 g are excreted as urea, 2.871 g are
excreted as ammonia, and 2.35 g of unassimilated
nitrogen plus 0.8 g of assimilated nitrogen are ex-
creted as feces.

C7 H12 O4 N + aO2 = bC6 H8 O2 N + cC6 H11 O6 N + dCH4 ON + eNH3 specific shrimp oxygen consumption rate as a function of the shrimp
size were analysed. For this, the statistical analysis used the Pearson
+ fH2 O + gCO2 (2)
correlation coefficient (r), as proposed by Santana Neto (2017).

2.4. Model calibration

3. Results and discussion
The proposed model was validated by comparing the specific rate of
feed consumption μr, ammonia nitrogen production µN _ NH3 , and oxygen 3.1. Elemental composition
consumption µO2 shr to values in the literature. For this, the shrimp
specific growth rate μshr was used from data of real cultivation and the The results obtained from the elemental analysis (%) and the
yield factors of shrimp biomass per feed Yshr/r, ammonia nitrogen per minimum formula (MF) elemental composition of feed, shrimp biomass,
feed YN _ NH3/ r , and oxygen consumption per feed YO2 shr / r , obtained from and feces are presented in Table 1.
the stoichiometric model. According to our chemical analysis, the nitrogen content of the feed
Shrimp mass over time was obtained from the experimental work of (Table 1) was determined as 6.0%, which is 0.4% higher than indicated
Nunes (2011), in which the moisture content was 78.75% as obtained by the manufacturer (5.6%). Shrimp biomass shows a much higher
by elemental analysis. Thus, dry mass (Mc in grams) was fitted at a time content of nitrogen (11.0%) compared to the feed. It is because the
(t in days) function by a monotone second-degree polynomial equation, animal incorporates nitrogen into biomass over time, forming its
resulting in Eq. (3) with R2 = 1.0000. Shrimp growth rate (Vc) was muscle fibers, for example. This process is inherent in the animal
obtained by the derivative of Eq. (3) with respect to time, resulting in growth. The elemental composition of feces shown in Table 1 represents
Eq. (4). a mixture of digested and undigested compounds, and a small amount
of ammonia, since shrimp excretion occurs mainly by the gills
Mc = 8.45 10 5 t2 + 9.91 10 5 t + 0.217 (3)
(Regnault, 1987; Schmidt-Nielsen, 2002).
Vc = dMc/ dt = 1.64 10 5 t + 9.91 10 5 (4)
From the ratio Vc/Mc we determined the specific shrimp growth μshr 3.2. Stoichiometric balance
(1/d). Through conversion yield factors Yshr/r (gshr/ gfeed), YN _ NH3/ r
(gN NH3/ gfeed), and YO2 shr / r (gO2/ gfeed ) , we estimated the specific rate of The coefficients resulting from the elemental balance, as shown in
feed consumption μr (gfeed/ gshrd), ammonia nitrogen generation µN NH3 Fig. 1, applied to the stoichiometric equation (Eq. (2)) is presented in
(gN NH3/ gshr d), and oxygen consumption µO2 shr (gO2 / gshr d) according Eq. (8).
to Eqs. (5), 6 and 7, respectively.
Table 1
µr = µshr / Yshr / r (5)
Feed, shrimp biomass and feces elemental composition (%) and their minimum
= µshr (YN _ NH3/ r / Yshr / r ) formula elemental composition in whole-number molar count (MF).
µN NH3 (6)
Element Feed¹ Shrimp biomass ² Feces³
µO2 shr = µshr (YO2 shr / r /Yshr / r ) (7) % MF % MF % MF

C 40.0 7 37.0 4 22.0 6

H 6.0 12 6.0 8 3.08 11
2.7. Statistical analysis
O 33.0 4 30.0 2 30.6 6
N 6.0 1 11.0 1 4.6 1
The specific growth rate as a function of the shrimp mass, the spe-
cific feed consumption rate as a function of the shrimp size and the 1, 2, 3
Values based on dry weight.

3
M.E.S. de Melo Filho, et al. Ecological Modelling 437 (2020) 109291

C7 H12 O4 N + 4.31O2 0.1231C6 H8 O2 N + 0.4831C6 H11 O6 N

+ 0.0197CH4 ON + 0.3544NH3 + 2.28H2 O
+ 3.59CO2 (8)

The stoichiometric model (Eq. (8)) proposes the nitrogen budget of

L. vannamei metabolism, considering the mass conservation principles
(Fig. 1). Alternate formulation and digestibility of feed used in aqua-
culture may change the stoichiometric equation coefficients, that can be
estimated for this new setup in growth condition using the proposed
methodology. This method can be extended to other species where the
digestibility of the feed and the chemical content of the feed, biomass,
and feces are known. From this stoichiometric equation, it is possible to
predict the mass quantity of products generated and obtain a global
mass balance for the system. Also, it supports establishing a control
system for shrimp farming. Mathematical modeling has been used
successfully in animal rearing operations, providing a quantitative un-
derstanding of cultivation-scale metabolic processes and aiding pro-
blem solving in real world applications (Chen and Malone, 1992; Sodré,
2007; Seginer, 2016; Fontes et al., 2019; Cheng et al., 2020; De Melo
Filho et al., 2020).

3.3. Specific rates

The Fig. 2 shows the specific growth rate μshr (1/d) of the shrimp
obtained in this work as a function of the total mass of the shrimp
(shrimp size in terms of wet weight). The data were estimated from
adjusted function to experimental data of shrimp growth and its deri-
vative Eqs. (3) and (4). A power law equation of μshr values fitted to the
actual total shrimp wet mass (W) resulted in Eq. (9) with R2 = 0.9999.

µshr = (4.39 10 2) W ( 5.38 t )

(9)
Fig. 3. (A) Specific rate of feed consumption μr and (B) ammonia nitrogen
From the stoichiometric model presented in Eq. (8), we estimated production µN _ NH3 , as a function of shrimp size in terms of wet weight.
the mass yield factors: Yshr / r = 0.0726 (gshr/ gfeed), and YN _ NH3/ r = 0.02851 Comparative plot between data from this work (continuous line) and estimated
(gN NH3/ gfeed), and YO2 shr / r = 0.7926 (gO2/ gfeed ) . based on literature methods Nunes and Parsons (2000) and Regnault (1987)
The specific rate of feed consumption, μr, and ammonia nitrogen (dashed line).
production, µN NH3 , were estimated as a function of W using these yield
factors Eq. (9), 5 and (6). The Fig. 3 shows both specific rates plotted
against shrimp wet mass, as well as data from the literature, which were
used to estimate the Pearson's correlation. Experimental values of μr
were obtained according to Nunes and Parsons (2000), and µN NH3
values were based on the average value of excretion of ammonia ni-
trogen (65%) described by Regnault (1987). The results of Pearson's
analysis showed r values between the model proposed in the present
work and values from the literature of 0.9995 and 0.9995 to μr and
µN NH3 , respectively.
Shrimp specific respiration rate, estimated by Eq. (7) and converted
to endogenous condition as a function of the shrimp wet mass, de-
termined in this work, is presented in Fig. 4 as well as the experimental

Fig. 4. Oxygen specific consumption rate of L. vannamei under endogenous

respiration condition as a function of shrimp size in terms of wet weight. The
continuous line represents values estimated by the model proposed in present
work. The dashed line represents an extrapolation of the model presented in
this study to literature data: circles) experimental data from Ponce-
Palafox et al. (1997b), squares) experimental data from Valenzuela-
Quiñónez et al. (2011).

data found in the literature for it (Ponce-Palafox et al., 1997b;

Fig. 2. Shrimp specific growth rate as the function of shrimp mass: experi-
mental data (dots) and adjusted power law equation (continuous line).
Valenzuela-Quiñónez et al., 2011). Endogenous specific respiration rate
*
µO2 shr was determined through data obtained by Rosas et al. (2001), in
which it was observed that shrimp under endogenous respiration pre-
sents around 49% lower values of rate when compared to shrimp with
the full digestive tract.

4
M.E.S. de Melo Filho, et al.
Pearson's analysis showed r values of 0.9998 and 0.9999 when
compared to Ponce-Palafox et al. (1997b) and Valenzuela-
Quiñónez et al. (2011) data, respectively, showing good agreement
between the model proposed and experimental data reported in the
literature.
In aquaculture the average specific growth rate (0.0199 (1/d),
Fig. 2, is used to compare cultures or experiments under different
conditions. However, the specific growth rate at a cultivation condition
decreases with increasing mass, as can be seen by the experimental data
and identified by the model proposed in this work. Assigning the mean
value of the specific growth rate to the whole cultivation will generate
underestimated and overestimated values according to the animal mass.
Also, all process control will be impaired since all other specific rates
are proportional to the specific growth rate.
4. Conclusion
The proposed model, using a stoichiometric equation based on the
conservation of elementary nitrogen, allows calculating both specific
consumption rate of feed and oxygen at different shrimp growth stages.
Also, it is possible to obtain from the equations ammonia, urea, feces,
undigested feed, and carbon dioxide kinetics, representing this rela-
tively complex system. Therefore, this work presents a tool for better
management in shrimp farming based on the environmental conditions
proposed in the present study, controlled and clear water systems, in-
cluding the prediction of required dissolved oxygen concentration and
feed amount at each growth phase, inlaying in reducing costs and en-
vironmental impact associated.
CRediT authorship contribution statement
Marcos Estevão Santiago de Melo Filho: Conceptualization,
Investigation, Methodology, Data curation, Writing. Marco Shizuo
Owatari: Data curation, Formal analysis, Writing - original draft,
Writing. José Luiz Pedreira Mouriño: Investigation, Methodology.
Bruno Augusto Mattar Carciofi: Investigation, Methodology. Hugo
Moreira Soares: Resources, Methodology.
Declaration of Competing Interest
The authors declare that they have no conflict of interest.
Acknowledgements
The authors acknowledge the National Council for Scientific and
Technological Development (CNPq) and Coordination for the
Improvement of Higher Education Personnel (CAPES) for the financial
support and M.E.S. Melo Filho scholarship, for research grant and fi-
nancial support to J.L.P. Mouriño (CNPq 308292 / 2014–6) and H.M.
Soares (CNPq Lattes: 2713109621898643); Chemical Engineering
Department (UFSC) and Marine Shrimp Laboratory (LCM - UFSC).
References
ASTM, 2012. D5373-08 standard test methods for instrumental determination of carbon,
hydrogen, and nitrogen in laboratory samples of coal. Annu. B. ASTM Stand. 5, 1–9.
Byron, C., Link, J., Costa-Pierce, B., Bengtson, D., 2011. Calculating ecological carrying
capacity of shellfish aquaculture using mass-balance modeling: narragansett Bay,
Rhode Island. Ecol. Model. 222 (10), 1743–1755.
Bett, C., Vinatea, L., 2009. Combined effect of body weight, temperature and salinity on
shrimp Litopenaeus vannamei oxygen consumption rate. Brazilian J. Oceanogr. 57,
305–314. https://doi.org/10.1590/S1679-87592009000400005.
Boyd, C.E., Teichert-Coddington, D., 1995. Dry matter, ash, and elemental composition of
pond-cultured Penaeus vannamei and P. stylirostris. J. World Aquac. Soc. 26, 88–92.
https://doi.org/10.1111/j.1749-7345.1995.tb00214.x.
Buglione-Neto, C., Mouriño, J.L., do Nascimento Vieira, F., da Silva, B.C., Jatobá, A.,
Seiffert, W., Andreatta, E., 2013. Métodos para determinação da digestibilidade
aparente de dietas para camarão marinho suplementadas com probiótico. Pesqui.
Agropecu. Bras. 48 (8), 1021–1027.
5
Ecological Modelling 437 (2020) 109291
Chen, S., Malone, R.F., 1992. A mathematical model for soft-shell crawfish production.
Aquacult. Eng. 11 (4), 217–229.
Cheng, W., Sun, Z., Liang, S., Liu, B., 2020. Numerical model of an aquaculture structure
under oscillatory flow. Aquacult. Eng., 102054.
De Melo Filho, M.E.S., Owatari, M.S., Mouriño, J.L.P., Lapa, K.R., Soares, H.M., 2020.
Application of nitrification and denitrification processes in a direct water reuse
system for Pacific white shrimp farmed in biofloc system. Aquacult. Eng., 102043.
Ebeling, J.M., Timmons, M.B., Bisogni, J.J., 2006. Engineering analysis of the stoichio-
metry of photoautotrophic, autotrophic, and heterotrophic removal of ammonia-ni-
trogen in aquaculture systems. Aquaculture 257, 346–358. https://doi.org/10.1016/
j.aquaculture.2006.03.019.
FAO, 2018. The State of World Fisheries and Aquaculture 2018 - Meeting the Sustainable
Development Goals. Rome.
FAO, 2015. FAO Statistical Pocketbook 2015. Food and Agriculture Organization of the
United Nations 978-92-5-108802-9.
FAO, 2014. FAO Statistics [WWW Document]. Food Agric. Organ. United Nations URL.
http://faostat3.fao.org/home/E%5Cnhttp://faostat3.fao.org/faostatgateway/go/to/
download/Q/QC/E%5Cnhttp://faostat3.fao.org/.
Fontes, R.F.C., Fey, J.D., de Oliveira, A.J.F.C., Barbieri, E., 2019. Numerical modeling as
supporting tool for aquaculture of oysters in a subtropical estuarine ecosystem. Bol.
Inst. Pesca. 45 (4).
Gao, W., Tian, L., Huang, T., Yao, M., Hu, W., Xu, Q., 2016. Effect of salinity on the
growth performance, osmolarity and metabolism-related gene expression in white
shrimp Litopenaeus vannamei. Aquac. Reports 4, 125–129. https://doi.org/10.1016/j.
aqrep.2016.09.001.
Hernández, M., Bückle, F., Palacios, E., Barón, B., 2006. Preferential behavior of white
shrimp Litopenaeus vannamei (Boone 1931) by progressive temperature-salinity si-
multaneous interaction. J. Therm. Biol. 31, 565–572. https://doi.org/10.1016/j.
jtherbio.2006.05.008.
Hopkins, J.S., Devoe, M.R., Holland, A.F., 1995. Environmental impacts of shrimp
farming with special reference to the situation in the continental United States.
Estuaries 18, 25–42. https://doi.org/10.2307/1352281.
Jayasankar, V., Jasmani, S., Nomura, T., Nohara, S., Huong, D.T.T., Wilder, M.N., 2009.
Low salinity rearing of the Pacific white shrimp Litopenaeus vannamei: acclimation,
survival and growth of postlarvae and juveniles. Japan Agric. Res. Q. 43, 345–350.
https://doi.org/10.6090/jarq.43.345.
Klemetson, S.L., Rogers, G.L., 1985. Aquaculture pond temperature modeling. Aquacult.
Eng. 4 (3), 191–208.
Lee, P.G., Lawrence, A.L., 1985. Effects of diet and size on growth, feed digestibility and
digestive enzyme activities of the marine shrimp, Penaeus setiferus Linnaeus. J. World
Maric. Soc. 16, 275–287. https://doi.org/10.1111/j.1749-7345.1985.tb00210.x.
Maicá, P.F., Borba, M.R.de, Martins, T.G., Wasielesky Junior, W., Maicá, P.F., Borba,
M.R.de, Martins, T.G., Wasielesky Junior, W., 2014. Effect of salinity on performance
and body composition of Pacific white shrimp juveniles reared in a super-intensive
system. Rev. Bras. Zootec. 43, 343–350. https://doi.org/10.1590/S1516-
35982014000700001.
Mente, E., 2003. Nutrition, physiology, and Metabolism of Crustaceans. Science
Publishers.
Mente, E., Coutteau, P., Houlihan, D., Davidson, I., Sorgeloos, P., 2002. Protein turnover,
amino acid profile and amino acid flux in juvenile shrimp Litopenaeus vannamei: ef-
fects of dietary protein source. J. Exp. Biol. 205, 3107–3122.
Nunes, A.J.P.2011. Noções sobre a elaboração de tabelas de alimentação para camarões
marinhos [www Document]. URLhttp://abccam.com.br/site/wp-content/uploads/
2011/02/Programa_Alimentar.pdf (accessed 10.01.2020).
Nunes, A.J.P., Parsons, G.J., 2000. Size-related feeding and gastric evacuation measure-
ments for the Southern brown shrimp Penaeus subtilis. Aquaculture 187, 133–151.
https://doi.org/10.1016/S0044-8486(99)00386-5.
Panchang, V., Cheng, G., Newell, C., 1997. Modeling hydrodynamics and aquaculture
waste transport in coastal Maine. Estuaries 20 (1), 14–41.
Paz, P.E., Roy, L.A., Davis, D.A., Quintero, H.E., 2011. Survival of post-larval Litopenaeus
vannamei following acclimation to low salinity waters at different temperatures. J.
World Aquac. Soc. 42, 575–579.
Ponce-Palafox, J., Martinez-Palacios, C.A., Ross, L.G., 1997a. The effects of salinity and
temperature on the growth and survival rates of juvenile white shrimp, Penaeus
vannamei, Boone, 1931. Aquaculture 157, 107–115.
Ponce-Palafox, J., Martinez-Palacios, C.A., Ross, L.G., 1997b. The effects of salinity and
temperature on the growth and survival rates of juvenile white shrimp, Penaeus
vannamei, Boone, 1931. Aquaculture 157, 107–115.
Regnault, M., 1987. Nitrogen excretion in marine and fresh water crustacea. Biol. Rev.
https://doi.org/10.1111/j.1469-185X.1987.tb00623.x.
Rosas, C., López, N., Mercado, P., Martínez, E., 2001. Effect of salinity acclimation on
oxygen consumption of juveniles of the white shrimp Litopenaeus vannamei. J.
Crustac. Biol. 21, 912–922.
Santana Neto, J.P.D.2017. Desenvolvimento de uma ferramenta computacional baseada
no método de lattice Boltzmann para a investigação da acústica interna de dutos.
Saoud, I., Davis, D., Rouse, D., 2003. Suitability studies of inland well waters for
Litopenaeus vannamei culture. Aquaculture 217, 373–383.
Schmidt-Nielsen, K., 2002. Animal Physiology: Adaptation and Environment. Santos
Press, São Paulo.
Seginer, I., 2016. Growth models of gilthead sea bream (Sparus aurata L.) for aquaculture:
a review. Aquacult. Eng. 70, 15–32.
Seidman, E.R., Lawrence, A.L., 2009. Growth. Feed digestibility, and proximate body
composition of juvenile Penaeus vannamei and Penaeus monodon grown at different
dissolved oxygen levels. J. World Maric. Soc. 16, 333–346.
Sodré, U., 2007. Modelos Matemáticos. UEL, Londrina.
Teleken, J.T., Galvão, A.C., Robazza, W.D.S., 2017. Comparing non-linear mathematical
M.E.S. de Melo Filho, et al.
models to describe growth of different animals. Acta Scientiarum. Anim. Sci. 39 (1),
73–81.
Timmons, M.B., Vinci, B.J., Davenport, M.T., Crum, M.K., 2001. A mathematical model of
low-head oxygenators. Aquacult. Eng. 24 (4), 257–277.
Timmons, M.B., Ebeling, J.M.2007. Recirculating aquaculture.
Ulanowicz, R.E., Platt, T., 1985. Ecosystem Theory For Biological oceanography:
Proceedings of a Symposium Sponsored By SCOR, NSERC, NSF, Unesco, and the US
Office of Naval Research Held At Laval University, Ste. Foy, Quebec, March 16-23,
1984 213 Canadian Government Pub Centre.
Valenzuela-Quiñónez, W., Rodríguez-Quiroz, G., Ponce-Palafox, J.T., Esparza-Leal, H.M.,
2011. Efecto de diferentes combinaciones de temperatura y salinidad sobre el
Ecological Modelling 437 (2020) 109291
consumo específico de oxígeno en el camarón blanco Litopenaeus vannamei. Rev. Biol.
Mar. Oceanogr. 46, 303–311.
Velasco, M., Lawrence, A.L., Castille, F.L., 1999. Effect of variations in daily feeding
frequency and ration size on growth of shrimp, Litopenaeus vannamei (Boone), in zero
- water exchange culture. Aquaculture 179, 141–148. https://doi.org/10.1016/
S0044-8486(99)00158-1.
Vinatea, L., 1997. Princípios Químicos De Qualidade Da água em aqüicultura: Uma
Revisão Para Peixes e Camarões. UFSC, Florianópolis.
Zotin, A.A., 2015. The united equation of animal growth. Amer. J. Life Sci. 3 (5),
345–351.