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Abstract

Juveniles (3.5 ± 0.3 g) of the white shrimp Litopenaeus vannamei were grown during 40 days with no water exchanges, no food addition and four initial densities (25, 50, 75 and 100 g m -3 , corresponding to between 8 and 32 shrimp m -2), to determine growth rates, which could be achieved using the periphyton growing on artificial substrates as the only food source. The experimental culture units were 12 polyethylene 1 m 3 cylindrical tanks with 4.8 m 2 of total submerged surface (bottom and walls), provided with 7.2 m 2 of artificial substrate (Aquamats™). There were no significant differences in the ammonia and nitrite concentrations determined in the four treatments (0.17-0.19 and 0.10-0.11 mg L -1 , respectively), which remained below the respective levels of concern for shrimp cultures. Mean survival was similar, and ranged from close to 91 to 97%, whereas there were significant differences in mean individual weight, which ranged from 11.9-10.6 g shrimp -1 for the two low initial densities (25 y 50 g m -3), to 8.3-7.7 g shrimp -1 for the other treatments. However, because of the high survival and of the higher initial density, the best biomass yield was with 100 g m -3 . The final nitrogen contents of sediment and water were lower than the initial values, and between 36 and 60% of the difference was converted into shrimp biomass. Cultivo de camarón blanco (Litopenaeus vannamei Boone, 1931) sin recambio de agua y sin adición de alimento formulado: un sistema amigable con el ambiente
Culture of Litopenaeus vannamei with no food addition
441
Lat. Am. J. Aquat. Res., 40(2): 441-447, 2012
DOI: 10.3856/vol40-issue2-fulltext-19
Research Article
Culture of white shrimp (Litopenaeus vannamei Boone, 1931) with zero
water exchange and no food addition: an eco-friendly approach
Juan Manuel Audelo-Naranjo1, Domenico Voltolina2 & Emilio Romero-Beltrán3
1Universidad Autónoma de Sinaloa, Facultad de Ciencias del Mar
Mazatlán, Sinaloa, México
2Centro de Investigaciones Biológicas del Noroeste, Laboratorio de Estudios Ambientales
UAS-CIBNOR, Mazatlán, Sinaloa, México
3Instituto Nacional de Pesca, Centro Regional de Investigación Pesquera
Mazatlán, Sinaloa, México
ABSTRACT. Juveniles (3.5 ± 0.3 g) of the white shrimp Litopenaeus vannamei were grown during 40 days
with no water exchanges, no food addition and four initial densities (25, 50, 75 and 100 g m-3, corresponding
to between 8 and 32 shrimp m-2), to determine growth rates, which could be achieved using the periphyton
growing on artificial substrates as the only food source. The experimental culture units were 12 polyethylene 1
m3 cylindrical tanks with 4.8 m2 of total submerged surface (bottom and walls), provided with 7.2 m2 of
artificial substrate (Aquamats™). There were no significant differences in the ammonia and nitrite
concentrations determined in the four treatments (0.17-0.19 and 0.10-0.11 mg L-1, respectively), which
remained below the respective levels of concern for shrimp cultures. Mean survival was similar, and ranged
from close to 91 to 97%, whereas there were significant differences in mean individual weight, which ranged
from 11.9-10.6 g shrimp-1 for the two low initial densities (25 y 50 g m-3), to 8.3-7.7 g shrimp-1 for the other
treatments. However, because of the high survival and of the higher initial density, the best biomass yield was
with 100 g m-3. The final nitrogen contents of sediment and water were lower than the initial values, and
between 36 and 60% of the difference was converted into shrimp biomass.
Keywords: Litopenaeus vannamei, artificial substrates, nutrient recycling, biofilm, nitrogen budget, water
quality, Mexico.
Cultivo de camarón blanco (Litopenaeus vannamei Boone, 1931) sin recambio de
agua y sin adición de alimento formulado: un sistema amigable con el ambiente
RESUMEN. Durante 40 días se cultivaron juveniles de camarón blanco Litopenaeus vannamei con un peso
individual de 3,5 ± 0,3 g y biomasas iniciales de 25, 50, 75 y 100 g m-3 (equivalente a 8-32 ind m-2), sin
cambios de agua y adición de alimento, para determinar la tasa de crecimiento usando como única fuente de
alimentación el perifiton desarrollado en sustratos artificiales. Se utilizaron estanques cilíndricos de polietileno
de 1 m3 con tres réplicas por tratamiento, con una superficie de 4,8 m2 (paredes y fondo) y 7,1 m2 de sustrato
artificial (Aquamats™). No se encontraron diferencias significativas entre las concentraciones de amonio
(0,17-0,19 mg L-1) y nitrito (0,10-0,11 mg L-1) determinadas en los cuatro tratamientos. La supervivencia fue
similar, variando entre 91 y 97%. La ganancia en peso individual fue significativamente mayor en los
tratamientos con menor biomasa inicial (25 y 50 g m-3), aunque por la mayor densidad inicial, el mejor
rendimiento en biomasa se observó en los cultivos sembrados con 100 g m-3. Los contenidos de nitrógeno
determinados al final del experimento, en el agua y sedimento, fueron inferiores a los valores iniciales, y entre
el 36 y 60% de sus diferencias se recuperaron en biomasa de camarón.
Palabras clave: Litopenaeus vannamei, sustrato artificial, reciclamiento de nutrientes, biopelícula, balance de
nitrógeno, calidad de agua, México.
___________________
Corresponding author: Juan Manuel Audelo-Naranjo (jmaudelo@gmail.com)
Latin American Journal of Aquatic Research
442
INTRODUCTION
The high cost and the large amounts of formulated
feed may become an important limiting factor for
intensive shrimp culture. In addition, shrimp
metabolism and leaching of organic substances from
food and feces may cause poor water quality and pond
bottom deterioration (Burford & Williams, 2001;
Avnimelech & Ritvo, 2003), because of the inverse
relationship between food assimilation efficiency and
culture density (Martin et al., 1998; Zaki et al., 2004).
There are several solutions to the water quality
problem: the traditional way is an increase of water
exchange rates. However, this practice increases the
operating costs due to the high water and energy
consumption, and the lower retention time of nutrients
within the culture systems, that would otherwise be
available for biogeochemical recycling by bacteria and
phytoplankton, thereby increasing the availability of
natural food (Jackson et al., 2003; Crispim et al.,
2007). Other solutions involve the removal of
nutrients either outside or within the culture system. In
the first case, nutrients are removed through different
combinations of physical, chemical and biological
processes, such as mechanical filters, settling tanks,
ozonation or U.V. irradiation, and various types and
designs of biological filters (Timmons et al., 2002;
Hussenot, 2003; Gutierrez-Wing & Malone, 2006).
An alternative is the promotion of growth of
natural planktonic or benthic microbial and microalgal
communities (bioflocs and periphyton, respectively)
present in the pond environment, because their
utilization of nutrients through autotrophic and
heterotrophic processes accelerates the removal of
organic and inorganic wastes, thus improving water
quality; in addition their biomass can be used as a
source of food by the cultivated organisms (Azim et
al., 2002; Avnimelech, 2005).
Both techniques have shown to increase fish and
shrimp production in semi-intensive or intensive
experimental or commercial cultures (Browdy et al.,
2001; Keshavanath et al., 2001; Lopes-Thompson et
al., 2002; Van Dam et al., 2002; Avnimelech, 2007).
In particular, several studies have shown that the
bacterial and microalgal biofilm growing on natural or
artificial submerged substrates (periphyton) may be
used successfully as the main or sole food source for
several freshwater or brackish water fish species
(Ramesh et al., 1999; Azim et al., 2002, 2004; Jana et
al., 2004; Keshavanath et al., 2004). However,
available information for shrimp culture is limited to
some studies on intensive cultures, with formulated
feed as the main food source (Bratvold & Browdy,
2001; Otoshi et al., 2001; Domingos & Vinatea, 2008;
Audelo-Naranjo et al., 2011).
The aim of this work was to evaluate the growth
and production of juvenile Pacific white shrimp L.
vannamei, maintained at four initial stocking densities
in experimental cultures with artificial substrates, with
zero water exchange and no food addition.
MATERIALS AND METHODS
The experiment lasted 40 days, from July 2 to August
10, 2009, and was performed on the grounds of a
commercial farm close to the Urías Estuary, Mazatlán,
Sinaloa, NW Mexico (23°25’N, 106°22’W), which is
the source of seawater used by the farm for pond
filling and water exchanges.
Juveniles (3.5 ± 0.3 g) of L. vannamei obtained
from this farm were stocked in triplicate tanks, with
four initial stocking densities (25 ± 7.5; 50 ± 2.3; 75 ±
2.8 and 100 ± 2.5 g m-3: T25, T50, T75 and T100,
respectively), equivalent to 8, 16, 24 and 32 ind m-2.
The experimental units were 12 cylindrical heavy-duty
polyethylene tanks (1 m3, bottom surface: 1.1 m2 and
submerged walls: 3.7 m2). One week before the
experiment, each tank received a 10 cm-deep layer of
homogenized, untreated sediments of an intensive
shrimp farm, and was filled with 1 m3 of 300 µm-
filtered estuary water.
Since no formulated feed was supplied, shrimp fed
only on the periphyton growing on the tank walls and
on the 7.1 m2 (both sides) of the artificial substrates
(Aquamats®, Meridian Applied Technology Systems,
Calverton, Maryland, USA), which had been remained
submerged in the pond water to allow periphyton
growth during the previous 30 days and added in a
circular arrangement to each tank one day before the
experiment, at a distance of 10 cm from the tank wall
(Audelo-Naranjo et al., 2010).
Throughout the study period, the experimental
units were maintained with zero water exchange, but
water was added once per week to each tank (on
average 5% of the tank volume) to compensate the
water lost by evaporation. Continuous aeration was
supplied by a 1 HP blower (0.768 kW; Sweetwater
1HP, Aquatic Eco-Systems, Apopka, FL, USA) to
avoid thermal stratification and for renovation of the
water in contact with the submerged surfaces.
Temperature and dissolved oxygen concentrations
were measured twice daily (8:00 and 18:00 h), using
an air-calibrated YSI model 57 oxygen meter (YSI,
Yellow Springs, OH, USA). Salinity and pH were
measured at 14:00 h with an Atago S/Mill-E
refractometer (Atago, Tokyo, Japan) and a Hanna HI
Culture of Litopenaeus vannamei with no food addition
443
98150 field pH meter (Hanna Instruments,
Woonsocket, RI, USA).
During the first day of the experiment, triplicate
samples of the water used to fill the tanks were filtered
through Whatman GF/C filters to determine the
concentrations of dissolved inorganic (N-NO3-; N-
NO2-; N-NH4+) and organic N (DON) using traditional
colorimetric techniques (Strickland & Parsons, 1972).
The particulate organic N (PON) retained on the filters
was determined using the method described by Holm-
Hansen (1968). The same methods were applied to
obtain information on the dissolved and particulate N
added weekly to each unit with the water used to
replace water losses. Unionized NH3 was calculated as
in Spotte & Adams (1983).
The initial and final organic nitrogen content of the
sediment and of the accompanying micro- and
meiobiota were determined with the Kjeldahl method
(AOAC, 2005), in triplicate un-sieved samples
obtained from the center and sides of each tank,
during the first and during the final day of the
experiment. The same method was used to determine
the initial and final N content of shrimp, and of
triplicate samples of the periphyton, obtained by
scraping with a scalpel a known area of the substrate
(Audelo-Naranjo et al., 2010).
At the beginning and at the end of the experiment,
all organisms were counted and weighed individually.
Survival (S%) was calculated as S% = 100 (Nf Ni-1),
where Nf and Ni are the final and initial numbers of
shrimp. The mean individual initial and final weights
of the specimens of each unit were used to calculate
the mean daily growth rate (GR) as GR = (Wf-Wi) t-1,
where Wf and Wi are the final and initial wet weights
(g), respectively, and t is the duration (days) of the
experiment.
The nitrogen budget of each experimental unit was
calculated with the equation:
Wi + Bi + Pi + Si = Sf + Bf + Pf + Wf (Hopkins et al.,
1993)
where the inputs were: Wi = total N content (dissolved
and particulate) of the water used for tank filling and
weekly additions, initial N contents of the shrimp
biomass (Bi), of periphyton (Pi), and of the sediment
and accompanying micro- and meiobiota (Si).
Outputs: final N content of the sediment (Sf), of
the shrimp biomass harvested (Bf), of the periphyton
(Pf), and of the water discharged at the end of the
experiment (Wf). An additional output were the
shrimps escaped overnight from the experimental
units, which were collected the following morning,
weighed, frozen and analyzed separately.
The mean values of temperature, dissolved oxygen,
pH, salinity and dissolved nutrient concentrations
were compared using repeated measures ANOVA
tests, or the equivalent Friedman’s non-parametric test
when the data were not normal or homoscedastic
(Kolmogorov-Smirnov and Bartlett’s tests). The mean
values of final yields, survival, individual weights, and
growth rate were compared using one-way ANOVA
or Kruskall-Wallis tests, after arcsine square root
transformation in the case of final survival. In all
cases, the level of significance was P = 0.05 (Zar,
1996).
RESULTS
There were no significant differences between the
mean water characteristics: morning and afternoon
mean water temperatures and dissolved oxygen
concentrations ranged from 28.5 to 31.5 °C and
between 6.1 and 6.3 mg L-1, respectively. The mean
pH value was 8.4 in all treatments, and salinity varied
between 37.4 and 37.5 g L-1. Ammonia and nitrites
remained between 0.17-0.19 and 0.10-0.11 mg L-1,
respectively, and the mean values of calculated
unionized ammonia were below 0.03 mg L-1; nitrates
varied between 0.26 and 0.29 mg L-1.
The mean DON and PON concentrations
determined in the four treatments were similar, and
ranged from 0.78 to 0.79 and from 1.27 to 1.29 mg
L -1, respectively (Table 1).
Mean final survival varied between 90.7 and
97.3%, without differences between treatments. In the
case of mean final weights and daily growth rates,
there were no statistically significant differences
between the two low biomass treatments (25 g m-3:
11.9 ± 2.6 g and 0.20 ± 0.05 g day-1; 50 g m-3: 10.6 ±
1.2 g and 0.17 ± 0.02 g day-1, respectively). Both were
significantly higher than the high-density treatments
with final weights of 8.3 ± 0.3 and 7.7 ± 0.5 g, and
growth rates of 0.10-0.11 ± 0.01 g day-1.
There were also significant differences in mean
final yields: the lowest and the highest (73.6 ± 4.7 and
199.6 ± 19.9 g experimental unit) were those of the
tanks stocked with 25 and 100 g of initial biomass.
The tanks stocked with 50 and 75 g gave intermediate
values (145.5 ± 3.5 and 139.6 ± 16.6 g), and there was
no difference between these two treatments (Table 2).
Since no food was added, the major nitrogen input
to the experimental units was that contained in the
initial sediment, followed by that of the periphyton
(12.5 and 7.4 g m-3, respectively, composed mainly by
bacteria and algae, as well as ciliates, nematodes,
occasionally with copepods and amphipods, and few
Latin American Journal of Aquatic Research
444
Table 1. Mean values (± standard deviation) of daily water temperature (T°C) and dissolved oxygen concentrations (DO:
morning and afternoon readings, am and pm, respectively), pH and salinity (afternoon readings), and weekly nutrient
concentrations in the cultures of Litopenaeus vannamei with artificial substrate (Aquamats™) and increasing initial
biomass (25 to 100 g m-3).
Tabla 1. Valores medio (± deviación estándar) de los registros diarios de temperatura (T°C) y oxígeno disuelto, pH,
salinidad y concentración semanal de nutrientes por tratamiento y horario en los cultivos de Litopenaeus vannamei con
sustrato artificial (Aquamats™) e incremento de biomasa inicial (25 a 100 g m-3).
T25 T50 T75 T100
T°C am 28.5 ± 2.6a 28.5 ± 2.6a 28.7 ± 2.6a 28.7 ± 2.6a
T°C pm 31.1 ± 2.3a 31.2 ± 2.4a 31.3 ± 2.4a 31.5 ± 2.4a
DO am (mg L-1) 6.3 ± 1.1a 6.2 ± 1.2a 6.3 ± 1.1a 6.3 ± 1.2a
DO pm (mg L-1) 6.2 ± 1.2a 6.3 ± 1.2a 6.1 ± 1.1a 6.1 ± 1.2a
pH 8.4 ± 0.1a 8.4 ± 0.1a 8.4 ± 0.1a 8.4 ± 0.1a
Salinity (psu) 37.4 ± 1.1a 37.5 ± 1.1a 37.5 ± 1.1a 37.5 ± 1.1a
N-NH4+ (mg L-1) 0.18 ± 0.07a 0.18 ± 0.07a 0.17 ± 0.08a 0.19 ± 0.07a
N-NH3 (mg L-1) 0.02 ± 0.01a 0.02 ± 0.01a 0.02 ± 0.01a 0.03 ± 0.01a
N-NO2- (mg L-1) 0.10 ± 0.07a 0.11 ± 0.07a 0.10 ± 0.08a 0.10 ± 0.09a
N-NO3- (mg L-1) 0.27 ± 0.08a 0.29 ± 0.07a 0.26 ± 0.08a 0.27 ± 0.09a
DON (mg L-1) 0.78 ± 0.10a 0.78 ± 0.10a 0.79 ± 0.08a 0.79 ± 0.09a
PON (mg L-1) 1.28 ± 0.31a 1.27 ± 0.35a 1.27 ± 0.35a 1.29 ± 0.43a
The equal superscripts indicate lack of significant differences (one-way repeated
measures ANOVAs, P = 0.05). (DON: dissolved organic nitrogen, PON: particulate
organic nitrogen).
Table 2. Mean values (± standard deviation) of production variables in the cultures of the white shrimp Litopenaeus
vannamei with artificial substrate (Aquamats™) and increasing initial biomass (25 to 100 g m-3).
Tabla 2. Valores medio (± desviación estándar) de las variables de producción de los cultivos de camarón blanco
Litopenaeus vannamei con sustrato artificial (Aquamats™) e incremento de biomasa inicial (25 a 100 g m-3).
T25 T50 T75 T100
Final survival (%) 91.4 ± 7.8a 96.3 ± 0.2a 90.7 ± 6.4a 97.3 ± 2.3a
Initial weight (g) 3.5 ± 0.3 3.5 ± 0.3 3.5 ± 0.3 3.5 ± 0.3
Final weight (g) 11.9 ± 2.6b 10.6 ± 1.2b 8.3 ± 0.3a 7.7 ± 0.5a
Growth rate (g day-1) 0.20 ± 0.05b 0.17 ± 0.02b 0.11 ± 0.01a 0.10 ± 0.01a
Yield (g) 73.6 ± 4.7a 145.5 ± 3.5b 139.3 ± 16.7b 199.6 ± 19.9c
Different superscripts indicate significant difference between values in the same row (One-way ANOVAs, P = 0.05,
a<b<c).
polychaetes). The contribution of initial biomass
(nitrogen content: 3.68 ± 0.09%, wet weight) was
between 0.9 to 3.7 g m-3, depending on the initial
density; the water used to fill the tanks and for weekly
additions added 2.3 g of N to the mean total inputs.
Among outputs, the N contents of sediment (11.8
to 10.5 g m-3), periphyton (3.7 to 3.5 g m-3, both
decreasing with increasing initial shrimp biomass),
and water (1.6 increasing with biomass to 2.0 g m-3)
were lower than the initial values: sediment was the
main N compartment, and the values decreased with
increasing stocking density. The increased N content
of the shrimp biomass (in this case increasing from 2.7
to 7.2 g m-3, depending on stocking density)
represented between 34 and 60% of the difference
between the initial and final values of sediment, water
and periphyton.
The N content of the shrimp escaped from their
tank represented only 0.2 to 0.4 g m-3. Therefore,
between 2 and 3.2 g of N were missing from the
Culture of Litopenaeus vannamei with no food addition
445
Table 3. Nitrogen budgets in the culture of the white shrimp Litopenaeus vannamei with artificial substrate (Aquamats™)
and increasing initial biomass (25 to 100 g m-3). The values are mean ± standard deviations of the N inputs and outputs
(g).
Tabla 3. Balance de nitrógeno en el cultivo de camarón blanco Litopenaeus vannamei con sustrato artificial
(Aquamats™) e incremento de biomasa inicial (25 a 100 g m-3). Valores medio ± deviación estándar de los ingresos y
egresos de N (g).
T25 T50 T75 T100
Inputs
Sediment 12.5 ± 0.7a 12.4 ± 0.2a 12.5 ± 0.7a 12.5 ± 0.7a
Periphyton 7.4 ± 0.03a 7.4 ± 0.02a 7.4 ± 0.03a 7.4 ± 0.03a
Water 2.3 ± 0.2a 2.3 ± 0.2a 2.3 ± 0.2a 2.3 ± 0.2a
Initial biomass 0.9 ± 0.06a 1.9 ± 0.1b 2.7 ± 0.01c 3.7 ± 0.01d
Total 23.1 ± 0.16a 23.8 ± 0.12b 24.9 ± 0.15c 25.8 ± 0.17d
Outputs
Sediment 11.8 ± 0.1d 11.2 ± 0.1c 10.8 ± 0.1b 10.5 ± 0.1a
Periphyton 3.7 ± 0.1b 3.4 ± 0.2a,b 3.6 ± 0.4a,b 3.5 ± 0.1a
Water 1.6 ± 0.06a 1.9 ± 0.06b,c 1.8 ± 0.06b 2.0 ± 0.1c
Final biomass 2.7 ± 0.2a 5.2 ± 0.1b 5.0 ± 0.6b 7.2 ± 0.7c
Escaped shrimps 0.2 ± 0.01a 0.2 ± 0.01a 0.4 ± 0.01b 0.2 ± 0.01a
Total 20.1± 0.22a 21.8 ± 0.18b 21.6 ± 0.9b 23.3 ± 0.07c
Missing 3.0 ± 0.2b 2.0 ± 0.2a 3.2 ± 1.0a,b 2.5 ± 0.7a,b
Different superscripts indicate significant difference between values in the same row (One-
way ANOVAs, P = 0.05, a<b<c).
global budget (Table 3). Assuming similar growth and
grazing pressure as on the Aquamats, the biofilm on
the tank walls (3.7 m2), and on aeration lines and air
stones (approximately 0.8-1 m2) could not contain
more than 0.2-0.25 additional g. Therefore, the
remaining N was probably lost to the atmosphere,
either as molecular N produced through denitrification
or more probably, in view of the aeration provided, as
gaseous NH3, which represented on average 11% of
the total ammonia present in the tanks.
DISCUSSION
The mean temperature, dissolved oxygen, pH and
salinity values were well within the appropriate ranges
for L. vannamei culture (Treece, 2000). Salinity was
higher than the isosmotic point (approximately 20 g
L-1) which may have a negative effect on the energy
budget of the white shrimp (Valdez et al., 2008),
although values as high as 40 g L-1 does not seem to
affect significantly the growth rate of this species
(Ponce-Palafox et al., 1997).
Dissolved and unionized ammonia, as well as
nitrite and nitrate values remained below the
respective safety levels for shrimp culture (7.09, 0.13,
25.7, and 232 mg L-1, respectively) (Frías-Espericueta
et al., 1999; Tsai & Chen, 2002; Lin & Chen, 2003).
This confirms the importance of the microbiota
adhered to the submerged surfaces, which used the
dissolved and particulate nutrients present in the water
column and in the sediments for their growth and
development. Thus, the periphyton served to maintain
water quality and to provide food for the cultured
shrimp, and the low final nitrogen concentrations in
water and sediment prove that its metabolism was
adequate to prevent deterioration of the culture
environment.
Additionally, the weight gain of the shrimp
confirms the importance of the biota associated to
artificial substrates as a natural food source for farmed
organisms. This community forms a complex food
web, in which the consumers and detritivores use
autotrophic microorganisms as their food source, and
the dissolved organic and inorganic nitrogen produced
by their metabolism is recycled into new biomass by
the autotrophic microalgae-bacteria mats. In the
trophic structure of the experimental units, shrimp
were the top consumers, and therefore were the final
beneficiaries of the nutrient and energy flow of these
closed systems.
Latin American Journal of Aquatic Research
446
The best final individual weights (11.9 and 10.6 g),
were obtained with the intermediate initial stocking
densities (25 and 50 g of initial biomass, equivalent to
8 and 16 ind m-2, respectively), and the best biomass
gain was with the highest initial biomass, although the
individual weight gain was not as high as that obtained
with the two lower stocking densities. In all cases, the
regime of closed culture promoted nutrient recycling,
provided sufficient food for the cultured organisms,
eliminated the cost of formulated feed and water
exchanges and was, at the same time, environmentally
friendly because it minimized the environmental
impact of nutrient-loaded effluents.
Culture experiments with mesocosms should not
be used to calculate possible yields of full-scale
cultures or for cost/benefit analysis, but to verify the
feasibility of a different approach for large-scale
cultures. However, on the basis of these results,
shrimp may be grown successfully in closed cultures
using submerged substrates. Once the proper
biological load is established through additional work
at the pilot scale, periphyton may maintain water
quality and serve at the same time as the main or even
as the only source of food, with yields close or higher
than the <1000 kg ha-1 which, according to official
statistics (SAGARPA, 2010), is the mean yield of
most Mexican semi-intensive shrimp farms.
ACKNOWLEDGMENTS
Supported by PROFAPI2011/016 and CIBNOR
project AC0.38. V. Nuñez, O. Zamudio, B. Mejía and
J. Madero of the Academic Group ‘Shrimp and Fish
Culture’ helped with the field and analytical work.
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