Aquaculture Research, 2002, 33, 621±626
Effects of stocking density and feeding levels on
growth and feed efficiency of Nile tilapia (Oreochromis
niloticus L.) fry
Abdel-Fattah M El-Sayed
Department of Aridland Agriculture, College of Food Systems, United Arab Emirates University, Al Ain, United Arab
Emirates
Correspondence: Abdel-Fattah M El-Sayed, Department of Aridland Agriculture, College of Food Systems, United Arab Emirates University,
Al Ain, United Arab Emirates. E-mail: a.elsayed@uaeu.ac.ae
Abstract
The effects of stocking density and feeding levels on
larval survival, growth rates, feed utilization effi-
ciency and body composition of Nile tilapia (Oreo-
chromis niloticus L.) fry were investigated in two
consecutive experiments. In experiment 1, swimup
fry (0.016 g average body weight) were stocked in
20-L fibreglass tanks, in a closed, recirculating
indoor system, at five stocking densities (3, 5, 10,
15 and 20 fry L 1) and fed a larval test diet (40%
crude protein) to apparent satiation, three times a
day, for 40 days. Fish survival, percentage weight
gain and specific growth rate (% SGR) were nega-
tively correlated with stocking density. The best
performance was achieved at 3 fry L 1. However,
no significant differences in growth parameters were
found between 3 and 5 fry L 1. Body composition
was not significantly affected by stocking density. In
experiment 2, fry (0.016 g average body weight)
were stocked at 5 fry L 1, and fed a larval test diet
(40% crude protein) at six feeding levels (10%, 20%,
25%, 30% and 35% BW day 1) and to satiation,
three times a day for 40 days. Fish growth rates
and survival were extremely poor at 10% feeding
level, and improved significantly with increasing
feeding levels up to 30%, and levelled off with fur-
ther increase in feeding levels. On the contrary, feed
conversion ratio (FCR) and protein production value
(PPV) were negatively correlated with feeding level.
The lowest feeding level (10%) produced signifi-
cantly lower body lipid and higher protein and ash
than other feeding levels. The present results suggest
ß 2002 Blackwell Science Ltd
that the optimum stocking density and feeding level
of Nile tilapia fry are 5 fry L 1 and 30% per day
respectively.
Keywords: Nile tilapia, stocking density, feeding
levels
Introduction
Tilapia is the second most important farmed fish in
the world, after carps. Tilapia culture is being prac-
ticed in most of the tropical, subtropical and temper-
ate regions. Great attention has been paid to tilapia
culture in recent years. The Food & Agriculture
Organization of the United Nations (FAO 1999)
reported that the production of farmed tilapia has
risen at an annual rate of 12% since 1984. As a
result, this production has jumped from 308 234
metric tonnes (mt) in 1988 to 1099 268 mt in
1999 with a 356% increase (FAO 2001). During
the same period, the value of these fish has increased
from 383 to 1427 million US$ (FAO 2001).
Despite the great potential of tilapia culture,
shortage of fry production to meet the increased
global demands remains one of the main obstacles
limiting the expansion of intensive culture of these
fishes. Information regarding larval culture, espe-
cially the effects of stocking density and feeding
regimes on fish performance is limited, inconsistent
and sometimes controversial. For example, Dambo &
Rana (1992) found that the growth rates of Nile
621
Stocking density and feeding levels of Nile tilapia fry A-F M El-Sayed
tilapia (Oreochromis niloticus L.) fry were negatively
correlated with stocking density ranging from 2 to
20 fry L 1. They suggested 5±10 fry L 1 as optimal
stocking density. Similarly, Rana (1981) considered
8 fry L 1 as optimal stocking density of O. mossam-
bicus. On the contrary, Gall & Bakar (1999) reported
that body size of tilapia fry was not affected by stocking
densities ranging from 10 to 200 fry L 1 when water
flow was uniform. Furthermore, Macintosh & De
Silva (1984) found that the relationship between
the survival of O. mossambicus and O. niloticus
female  O. aureus male fry and stocking density
was not consistent.
Similar discrepancies were reported regarding the
effects of feeding levels on the performance of tilapia
fry. MeÂlard & Philippart (1980) (cited by Coche
1982) reported that feeding rates of 20±30% were
adequate for Nile tilapia weighing 0±5 g, while
Santiago, Aldaba & Reyes (1987) found that
30±45% BW day 1 was required for optimum per-
formance of Nile tilapia fry. On the other hand,
Macintosh & De Silva (1984) found that growth
and survival of O. mossambicus fry was improved
with increasing feeding level from 6% to 24%
BW day 1 irrespective of fish stocking density.
It is evident therefore that further studies are
needed to verify the effects of stocking density and
feeding regimes on tilapia fry under hatchery condi-
tions. The present study was conducted in the
Department of Aridland Agriculture, College of Food
Systems, United Arab Emirates University, Al-Ain,
UAE, through two consecutive experiments, to in-
vestigate the effects of stocking density and feeding
levels on the growth, survival, feed conversion ratio
(FCR) and body composition of Nile tilapia fry.
Materials and methods
Fish and culture facilities
Nile tilapia fry were produced from tilapia brood-
stock kept in captivity in the aquaculture unit,
College of Food Systems, United Arab Emirates
University, Al-Ain, UAE. Fertilized eggs were re-
moved from the mothers' mouths and stocked in
hatching jars provided with upward aeration to
keep stirring them. After hatching, fry were stocked
into 50-L glass aquaria until the yolk sac absorption
was completed. Fry were then fed a commercial diet
(40% crude protein) for 4 days.
The fry were stocked into 20-L fibreglass tanks in
a closed, recirculating indoor system. The tanks
622
Aquaculture Research, 2002, 33, 621±626
were provided with central drainage pipes sur-
rounded by outer sleeves pipes, perforated at the
bottom, to facilitate self-cleaning and waste re-
moval. The culture system was provided with a
biological filter, aeration through an air compressor
and thermostat to keep water temperature at 27  C.
About 15% of the water was replaced by new fresh-
water daily. Lighting in the culture unit was set at
14:10 L:D cycle. Water quality parameters includ-
ing dissolved oxygen (DO), ammonia (NH4-N), ni-
trites (NO2-N), nitrates (NO3-N) and pH were
monitored weekly.
Experimental design
Experiment 1
The first experiment was designed to study the
effects of stocking density on fish survival, growth
rates, feed utilization efficiency and body compos-
ition. Triplicate groups of swimup fry (0.016 g aver-
age weight) were stocked in the rearing tanks at five
stocking densities (3, 5, 10, 15 and 20 fry L 1). The
fish were fed an experimental diet (40% crude pro-
tein, 400 kcal GE 100 g 1) to apparent satiation,
three times a day (08:00, 12:00 and 17:00 h) for
40 days. Fish were weighed collectively at 10-day
intervals, and their average weights recorded.
Experiment 2
The second experiment was conducted to investigate
the effects of feeding levels on the performance of
Nile tilapia fry reared in the same culture system
and under the same conditions used in experiment 1.
Fish fry were stocked in triplicate tanks in the
rearing tanks at a density of 5 fish L 1 (the optimal
stocking density attained from experiment 1) and
fed the larval test diet (40% crude protein, 400
kcal GE 100 g 1) at six feeding levels (10, 20, 25,
30 and 35% BW day 1) and to apparent satiation,
three times a day (08:00, 12:00 and 17:00 h) for
40 days. Fish were weighed collectively at 10-day
intervals, and their average weights determined.
Body composition analysis
At the end of each experiment, all fry in each tank
were netted, weighed and their individual lengths
recorded, and finally frozen for final body compos-
ition analyses. Initial body analyses were performed
ß 2002 Blackwell Science Ltd, Aquaculture Research, 33, 621±626
Aquaculture Research, 2002, 33, 621±626 Stocking density and feeding levels of Nile tilapia fry A-F M El-Sayed

Table 1 Effects of stocking density on the performance of Nile tilapia (Oreochromis niloticus L.) fry in experiment 1

Density (fry L 1) Final weight (g fish 1) % gain SGR* Survival (%) FCR² PPV³

3 0.89a 5438a 10.05a 100a 3.45a 8.69a

5 0.83a 5063a 9.87a 100a 2.86b 10.99a
10 0.56b 3375b 8.89b 95b 2.74b 9.26a
15 0.41c 2438c 8.11bc 94b 2.65b 8.70a
20 0.36c 2125c 7.78c 90bc 2.77b 8.23a

*Specific growth rate (%) ˆ 100 (ln final weight ln initial weight)/time (days). ²Feed conversion ratio ˆ dry feed intake (g)/wet weight
gain (g). ³Protein production value (%) ˆ 100 (protein gain/protein fed). Values in the same column with different superscripts are
significantly different (P ˆ 0.05).

Weight (g/fish) to test for the differences among treatment means

1 when F-values from the anova were significant.
3 fish/I 5 fish/I 10 fish/I 15 fish/I 20 fish/I

0.8 Results and discussion

0.6
0.4
0.2
0
0 10 20
Time (Days)
Figure 1 Growth rates of Nile tilapia (Oreochromis niloticus
L.) fry reared at different stocking densities in experiment 1.
on a pooled sample of 200 fry, which were weighed
and frozen prior to the study. Proximate analyses
of whole body water, protein, lipid and ash were
performed according to standard AOAC (1995)
methods.
Statistical analyses
Fish growth rates, feed utilization efficiency and
body composition results obtained from both experi-
ments were subjected to one-way analyses of vari-
ance (anova). Orthogonal polynomial procedures
(Gill 1981) were used to compare means at
P ˆ 0.05. Least significant difference (LSD) was used
Experiment 1
The results of experiment 1 revealed significant
effects of stocking density on larval survival, growth
rates and feed utilization efficiency (Table 1, Fig. 1).
Fish survival was reasonably good at all stocking
densities and ranged from 90% to 100%. This find-
ing may indicate that stocking density might have a
limited effect on fish survival. Similar results have
been reported by Dambo & Rana (1992) who found
that survival of Nile tilapia fry was between 94.5%
30 40 and 100% at stocking densities ranging from 2 to 20
fry L 1. The relatively increased mortality at 20
fry L 1 (10% mortality, P , 0.05) was most likely
due to cannibalism (not stocking density) as has
been reported with other tilapia species including
O. mossambicus (Uchida & King 1962; Macintosh &
De Silva 1984), O. niloticus (Dambo & Rana 1992)
and tilapia hybrids (Macintosh & De Silva 1984).
These studies demonstrated that cannibalism could
be a main cause of tilapia fry mortality.
Specific growth rate (% SGR) and percentage
weight gain were negatively correlated (P , 0.05)
with stocking density. However, no significant dif-
ferences in growth parameters (P . 0.05) were
found between 3 and 5 fry L 1. Further increases
in stocking density resulted in significantly poor
growth rates (P , 0.05). This finding indicated
that 5 fry L 1 is optimal for hatchery reared Nile
tilapia fry. This result is in agreement with that of
Silva, Souza, Padua, Dalacorte & Goncalves (2000),
who found that the growth of tetra hybrid red tilapia
decreased with increasing stocking density.

ß 2002 Blackwell Science Ltd, Aquaculture Research, 33, 621±626 623

Stocking density and feeding levels of Nile tilapia fry A-F M El-Sayed Aquaculture Research, 2002, 33, 621±626

The negative correlation between growth rates
and stocking density of fish fry has been investigated
by a number of authors. Macintosh & De Silva
(1984) reported that increasing stocking density of
O. mossambicus fry might have lead to diminishing
social dominance, leading to higher survival but
lower individual growth rates. Similar results were
reported on Nile tilapia fry (Dambo & Rana 1992).
It has also been reported that increasing fish dens-
ity leads to `social stress' causing `chronic stress
response'. This leads, in turn, to impaired fish
growth, presumably due to the mobilization of diet-
ary energy by the physiological alterations provoked
by the stress response (Kebus, Collins, Brownfield,
Amundson, Kayes & Malison 1992). In support,
Barcellos, Nicolaiewsky, de Souza & Lulhier (1999)
found that the resting plasma cortisol concentra-
tions of Nile tilapia fingerlings increased with in-
creasing the stocking density indicating a chronic
stress response, attributable to social stress.
Furthermore, King, Howell, Huber & Bengtson
(2000) suggested that stress due space availability
was the primary factor inhibiting the growth of
Summer flounder (Paralichthys dentatus L.) larvae
stocked at high densities. Increasing stocking dens-
ity may also cause deterioration in water quality,
leading to stressful conditions (Barton & Iwama
1991; Pankhurst & Van der Kraak 1997). However,
in the present work, the high water exchange rate
(three times per h) and the continuous removal of the
wastes and metabolites may exclude this assumption.
On the other hand, the FCR was not significantly
affected (P . 0.05) by stocking density, except at 3
fry L 1, which showed significantly lower feed util-
ization efficiency than at other stocking densities.
This may have been due to the lack of competition
for food and/or social hierarchy at 3 fry L 1. In
addition, the difficulty of tracing feed particles may
have lead to the reduction of feed consumption, and
to the flush of uneaten feed with the drainage water,
causing the deterioration of feed utilization effi-
ciency. Meanwhile, Silva et al. (2000) found that
feed consumption of tetra hybrid red tilapia in-
creased with increasing stocking density. Body com-
position was not significantly affected by stocking
density (Table 2).

Table 2 Effects of stocking density on body composition of

Nile tilapia fry (% dry weight) in experiment 1
Experiment 2

Density The effects of feeding levels on larval performance in

(fry L 1) Water Protein Lipids Ash experiment 2 are given in Table 3. Fish survival, %
weight gain and SGR were significantly affected by
Initial 81.73 68.36 14.55 14.39 feeding levels (P , 0.05). Growth parameters and
3 77.88a 65.77a 18.93a 12.35a survival were extremely poor at the 10% level, and
5 76.50a 64.92a 21.28a 12.30a
significantly improved (P , 0.05) with increasing
10 78.83a 65.34a 21.03a 13.49a
15 78.76a 66.93a 19.86a 13.20a feeding levels up to 30% (Fig. 2). Further increases
20 78.23a 66.47a 20.30a 13.24a in feeding levels did not result in any improvement
in fish growth rates. On the other hand, FCR was
Values in the same column with different superscripts are signifi- better at the 10% feeding level than at other feeding
cantly Different (P ˆ 0.05). levels (P , 0.05).

Table 3 Effects of feeding levels on the performance of Nile tilapia fry in experiment 2

Feeding level (%) Final weight (g fish 1) % gain % SGR Survival (%) FCR PPV

10 0.26a 2067a 7.69a 61a 1.27a 11.70a

20 0.35b 2833b 8.43b 86b 1.98b 11.00a
25 0.41b 3333b 8.83b 85b 2.13b 10.49a
30 0.59c 4833c 9.74c 87b 2.20b 9.00a
35 0.58c 4750c 9.70c 82b 2.98c 9.82a
Satiation 0.62c 5083c 9.86c 96c 2.86c 9.23a

See footnotes of Table 1. Values in the same column with different superscripts are significantly different (P ˆ 0.05).

624 ß 2002 Blackwell Science Ltd, Aquaculture Research, 33, 621±626

Aquaculture Research, 2002, 33, 621±626 Stocking density and feeding levels of Nile tilapia fry A-F M El-Sayed

Weight (g/fish)
have been due to the increased rate of feed passage
0.7
through the digestive tract and depressed feed digest-
10% 20% 25% 30% 35% Satiation
ibility and metabolism (Brett, Shelborn, & Shoop
0.6
1969; Bret & Groves 1979; Smith 1989 and Mazur,
Tillapaugh, Brett & Iwama 1993). Furthermore, at
0.5
higher feeding levels, significant amounts of feed
were flushed from the tanks through the drainage
0.4
pipes, leading to poor feed utilization efficiency, as
reported by Macintosh & De Silva (1984).
0.3
Except at the 10% feeding level, body composition
was not significantly affected by feeding regimes
0.2
(P . 0.05). The lowest feeding level (10%) pro-
duced significantly lower body lipids and higher
0.1
protein and ash contents than other feeding levels
(Table 4).
0
0 10 20 30 40
Time (days)

Figure 2 Growth rates of Nile tilapia fry fed a test diet at References
varying feeding levels in experiment 2.
Association of Official Analytical Chemists (AOAC) (1995)
Official Methods of Analysis. Association of Official
Table 4 Effects of feeding levels on body composition Analytical Chemists, Arlington, VA.
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