Aquaculture 231 (2004) 393 402

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Effects of photoperiod on the performance of

farmed Nile tilapia Oreochromis niloticus:
I. Growth, feed utilization efficiency and survival
of fry and fingerlings
Abdel-Fattah M. El-Sayed *, Mamdouh Kawanna
Department of Aridland Agriculture, College of Food Systems, United Arab Emirates University,
Al Ain, United Arab Emirates
Received 16 May 2003; received in revised form 5 November 2003; accepted 7 November 2003

Abstract
The effect of photoperiod on survival, growth rates and feed utilization efficiency of Nile
tilapia (Oreochromis niloticus L.) fry and fingerlings was investigated in two consecutive
experiments. In Experiment 1, triplicate groups of 75 swim-up fry (0.02 g) were stocked in 25-l
fiberglass tanks, in recirculating indoor system. The fish were exposed to four photoperiod
(light:dark, L:D) cycles (24L:0D, 18L:6D, 12L:12D and 6L:18D). Light intensity was kept
constant at 2500 lx throughout the study. The fish were fed a tilapia diet (35% crude protein, 350
kcal GE/100 g) at a daily rate of 30 20% BW, three times a day for 60 days. The best weight
gain, specific growth rate (SGR), feed efficiency and fish survival were achieved at 24L:0D and
18L:6D, without significant differences between them. Fry performance was significantly retarded
by reducing light phase (12L:12D and 6L:18D). In the second experiment, triplicate groups of 40
fingerlings (mixed sexes) (2.4 F 0.05 g) were stocked in 0.4-m3 fiberglass tanks and exposed to
the same light intensity and photoperiod cycles used in Experiment 1. The fish were also fed the
same diet used in Experiment 1, at 5 4% BW/day, three times a day for 90 days. The fish
performance was not significantly affected by photoperiods. These results revealed that Nile tilapia
fry, but not fingerlings, reared in indoor, recirculating systems are significantly affected by
photoperiod. The insignificant difference in fry performance between 24L:0D and 18L:6D groups
suggests that a 18L:6D cycle be used in case of larval rearing, while shorter light phases are

* Corresponding author.
E-mail address: a.elsayed@uaeu.ac.ae (A.-F.M. El-Sayed).
0044-8486/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2003.11.012

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A.-F.M. El-Sayed, M. Kawanna / Aquaculture 231 (2004) 393402

suggested for optimal growth, feed efficiency and survival of fish fingerlings, taking into
consideration the cost of electricity.
D 2004 Elsevier B.V. All rights reserved.
Keywords: Photoperiods; Nile tilapia; Fry; Fingerlings; Growth; Feed efficiency; Survival

1. Introduction
Tilapia are the third most important cultured fish group in the world, after carps and
salmonids (FAO, 2002). Tilapia culture is also one of the fastest growing farming activities,
with an average annual growth rate of 13.4% during 1970 1999 (FAO, 2002). As a result,
the production of farmed tilapia has increased from 383,654 mt in 1990 to 1,265,780 mt in
2000 (FAO, 2002). During the same period, the value of these fish has increased from
US$515.2 million to US$1706.5 million. Nile tilapia is by far the most important farmed
tilapia species in the world. The production of farmed Nile tilapia reached 1,045,100 mt in
2000, representing 82% of total production of farmed tilapia, with a value of US$1250.4
million (FAO, 2002).
The intensive culture of tilapia under controlled management systems is widely
expanding to meet the increasing demands for these fishes, especially in developing
countries. In this regard, the use of closed culture systems has received a considerable
attention, and is becoming more common worldwide, particularly in arid areas that face
shortage in fresh water or brackish water, or in areas where environmental parameters, such
as salinity and temperature, are outside the tolerance range of tilapia (Muir et al., 2000).
Tilapia can tolerate a wide range of water temperature, dissolved oxygen (DO), salinity,
pH, light intensity and photoperiods. However, the determination of optimal environmental
conditions for cultured tilapia in closed systems is essential for the maximization of its
production, profitability and sustainability (Muir et al., 2000). The effects of most of these
factors on tilapia in different culture systems have been extensively studied (Kirk, 1972;
Chervinski, 1982; Philippart and Ruwet, 1982). However, our knowledge on the effect of
photoperiod on tilapia in recirculating culture systems is limited.
Photoperiod acts as an artificial Zeitgeber (cue or synchronizer), regulating the daily
endogenous rhythms in fish (Duston and Saunders, 1990; Biswas et al., 2002) and also
affects fish growth, locomotor activity, metabolic rates, body pigmentation, sexual
maturation and reproduction (Gross et al., 1995; Silva-Garcia, 1996; Boeuf and Le Bail,
1999; Trippel and Neil, 2002; Biswas and Takeuchi, 2002; Biswas et al., 2002). On the
other hand, the growth and metabolic rates of several other species were not significantly
affected by photoperiods (Imsland et al., 1995; Hallaraker et al., 1995; Purchase et al.,
2000). Meanwhile, photoperiod may positively affect larval stages, but not juvenile stages
(Barlow et al., 1995).
Tilapia are generally diurnal feeders, feeding at different times of the day. Yousuf
Haroon et al. (1998) found that feeding activity of tilapia hybrids (O. mossambicus  O.
niloticus) reared in ponds was continuous during daytime, with a single feeding peak at
around afternoon-dusk. Similar results have been reported with the same fish reared in a

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395

closed system, where the maximum feeding activity occurred during the afternoon
(Zavyalov and Lavrovskii, 2001). In addition, Nile tilapia exhibit a regular diurnal feeding
rhythm with maximum feeding intensity between 5:00 and 8:00 a.m., and cease feeding
between 14:00 and 18:00 p.m. (Harbott, 1975). These studies clearly pointed out the role of
photoperiod on feeding activity and growth of these fishes. However, few studies have
considered the effect of photoperiod on tilapia growth, feed efficiency and metabolism,
physiological functions and reproduction under controlled culture conditions (Ross and
McKinney, 1988; Lourenco et al., 1998; Ridha and Cruz, 2000; Biswas and Takeuchi,
2002; Biswas et al., 2002).
We conducted a series of experiments to investigate the effect of photoperiod on the
performance of Nile tilapia (Oreochromis niloticus) reared intensively in a recirculating
system. The present study was carried out in two consecutive experiments to address the
effects of photoperiod on the growth, feed utilization efficiency and survival of fry and
fingerling fish.

2. Materials and methods

2.1. Fish and culture facilities
Nile tilapia fry and fingerlings were produced from tilapia broodstock kept in captivity
in the aquaculture facility, College of Food Systems, United Arab Emirates University, AlAin, UAE. The culture units consist of fiberglass tanks in a recirculating indoor system.
The tanks were provided with central drainage pipes surrounded by outer sleeves pipes,
perforated at the bottom, to facilitate self-cleaning and waste removal. The culture system
was also provided with a biological filter (Plastic tubing structures), continuous aeration
through an air compressor (Hick Hargreaves, UK) and heaters, with thermostats, to keep
water temperature at 27 jC. About 10% of the water was replaced daily by new fresh
water with the same temperature. Water quality parameters, including DO (Oxygen meter,
YSI, model 58), ammonia (NH4-N), nitrates (NO3-N) and nitrites (NO2-N) (Orion
Aquafast, Germany) and pH (pH meter, Jenway, UK) were monitored weekly. The
average values of these parameters throughout the study were: DO = 6.7 F 1.4 mg/l,
NH 4 -N = 0.053 F 0.002 mg/l, NO 3 -N = 11.4 F 1.32 mg/l, NO 2 V 0.05 mg/l and
pH = 8.1 F 0.06.
2.2. Experimental design
2.2.1. Experiment 1
The first experiment was designed to study the effect of photoperiod on growth rates,
feed utilization efficiency and survival of Nile tilapia fry. Triplicate groups of 75 swim-up
fry (0.02 g average weight) were stocked into 25-l rearing tanks (described in 2.1), with a
water flow rate of 1 l/m. The fish were exposed to four photoperiod (light:dark, L:D) cycles
(24L:0D, 18L:6D, 12L:12D and 6L:18D), using fluorescent lamps. Photoperiods were
controlled by a 24-h timer (Multi 9, Merlingerin, Germany). Light intensity was kept
constant at 2500 lx throughout the study.

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A.-F.M. El-Sayed, M. Kawanna / Aquaculture 231 (2004) 393402

The fish were fed an experimental diet (35% crude protein, 350 kcal GE/100 g) at a
daily rate of 30% BW/day, reduced to 20% BW/day at the beginning of the second
month (as recommended by El-Sayed, 2002), for 60 days. The diet was provided three
times a day at 8:00, 12:00 and 17:00 h, except for the 6L:18D group which was fed
twice a day at 8:00 and 13:00 h. All fish in each tank were weighed at 15-day
intervals, their average weights recorded, and the daily rations readjusted accordingly.
At the end of Experiment 1, all fish in each tank were netted, weighed and the average
final weights recorded. The undersized fingerlings were netted out and the rest of the
fish were transferred into 0.4-m3 fiberglass tanks (used in the second experiment)
having a water flow rate of 10 l/m. They were fed the test tilapia diet for a 10-day
acclimation period, to adapt them to the new tanks, stocking density and water flow
rate.
2.2.2. Experiment 2
The second experiment was conducted to investigate the effects of photoperiod on
survival, growth rates, feed utilization efficiency and survival of Nile tilapia fingerlings.
Forty fish of almost similar sizes were selected from each tank, weighed and their average
weight recorded (2.4 F 0.05 g). Triplicate groups of fish were stocked in the 0.4-m3 tanks
and exposed to the culture conditions, light intensity and photoperiod cycles used in
Experiment 1.
The fish were fed the same diet used in Experiment 1, for 90 days, at a daily rate of 5%
BW/day (reduced to 4% BW/day at the beginning of the third month), three times a day
(8:00, 13:00 and 17:00 h), except for the 6L:18D group where the diet was offered twice a
day at 8:00 and 13:00 h. All fish from each tank were weighed at 15-day intervals, their
average weights recorded, and daily rations readjusted accordingly. At the end of
Experiment 2, all fish in each tank were netted, weighed and their average final weight
recorded.
2.3. Statistical analyses
A one-way analysis of variance (ANOVA) was conducted to test the effect of
photoperiod on the growth rates, feed utilization efficiency and survival of fish fry and
fingerlings, using the computer program SPSS (SPSS Version 11.0.0, 2003). Least
significant difference (LSD) was used to compare means at P < 0.05, as described by Gill
(1981).

3. Results
3.1. Experiment 1
The results of Experiment 1 indicated that photoperiod significantly affected fish
survival and growth performance (Table 1 and Fig. 1). The long light phases (24 and 18
light hours) produced the best fish survival (89% and 85%), percentage weight gain (6050%
and 6100%), specific growth rate (% SGR) (6.87% and 6.88%) and feed conversion ratio

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397

Table 1
Effects of photoperiod on growth rates and feed utilization efficiency of Nile tilapia fry (mean F S.E.)
L:D cycle
24L:0D
18L:6D
12L:12D
6L:18D

IW1
0.02
0.02
0.02
0.02

FW2

% Gain3
a

1.23 F 0.08
1.24 F 0.09a
0.95 F 0.09b
0.65 F 0.07c

SGR4
a

6050 F 376
6100 F 425a
4650 F 449b
3150 F 344c

FCR5
a

6.87 F 0.16
6.88 F 0.12a
6.42 F 0.18b
5.80 F 0.19c

Survival (%)
a

1.52 F 0.12
1.55 F 0.07a
1.78 F 0.07ab
1.85 F 0.12b

89 F 3.60a
85 F 1.15ab
76 F 3.84b
78 F 6.49ab

Values in the same column with different superscripts are significantly different ( P = 0.05).
1
Average initial weight (g/fish).
2
Average final weight (g/fish).
3
100(FW IW/IW).
4
Specific growth rate = 100(ln FW ln IW)/time (days).
5
Feed conversion ratio = dry feed given (g)/wet weight gain (g).

(FCR) (1.52 and 1.55), respectively. However, the difference between these two photoperiods was not significant ( P>0.05). The growth and feed efficiency of fry were
significantly retarded by reducing light phase to 12 ( P < 0.01) and 6 ( P < 0.05) h.

Fig. 1. Effect of photoperiod on the growth of Nile tilapia fry.

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A.-F.M. El-Sayed, M. Kawanna / Aquaculture 231 (2004) 393402

Table 2
Effects of photoperiod on growth rates and feed utilization efficiency (mean F S.E.) of fingerling Nile tilapia
L:D cycle
24L:0D
18L:6D
12L:12D
6L:18D

IW1

FW2
a

2.33 F 0.09
2.43 F 0.07a
2.34 F 0.10a
2.44 F 0.06a

% Gain3
a

49.60 F 1.67
51.35 F 2.15a
46.80 F 2.71a
46.00 F 3.30a

SGR4
a

2024 F 24.0
2013 F 88.5a
1900 F 85.0a
1786 F 110.0a

FCR5
a

3.40 F 0.10
3.39 F 0.11a
3.33 F 0.05a
3.27 F 0.07a

Survival (%)
a

1.22 F 0.09
1.27 F 0.14a
1.32 F 0.08a
1.31 F 0.13a

95.0 F 0.00a
95.0 F 5.00a
95.0 F 2.50a
96.7 F 2.50a

Values in the same column with different superscripts are significantly different ( P = 0.05).
1
Average initial weight (g/fish).
2
Average final weight (g/fish).
3
100(FW IW/IW).
4
Specific growth rate = 100(ln FW ln IW)/time (days).
5
Feed conversion ratio = dry feed given (g)/wet weight gain (g).

3.2. Experiment 2
On the contrary to Experiment 1, the results of Experiment 2 revealed that fish survival,
percentage weight gain, SGR and FCR of mixed-sex fingerling tilapia were not significantly affected by photoperiod ( P>0.05) (Table 2).

4. Discussion
It has been suggested that freshwater fish species are more sensitive to photoperiod than
marine and diadromous species (Imsland et al., 1995). However, the response of marine
species to photoperiods has been well investigated, while less information is available on
freshwater species.
The results of the present study indicated that the response of Nile tilapia to photoperiod
cycles depends on fish developmental stage. Tilapia fry were more sensitive to photoperiod
than fingerlings and juveniles. Fish fry subjected to long light periods (24 and 18 h) had
significantly better growth and feed utilization efficiency than those exposed to intermediate or short light periods (12 or 6 h). Similar results have been reported with several
marine fish larvae, where the growth rates were improved with increasing light periods. The
growth of black porgy Mylio macrocephalus (Kiyono and Hirano, 1981), seabream Sparus
aurata (Tandler and Helps, 1985) and rabbitfish Siganus guttatus (Duray and Kohno, 1988)
larvae was best under continuous light, while the growth of sole Solea solea (Fuchs, 1978),
European seabass Dicentrarchus labrax (Barahona-Fernandes, 1979), barramundi Lates
calcarifer (Barlow et al., 1995) and snapper Pagrus auratus (Fielder et al., 2002) larvae was
better at 18 and 24 h light periods than at 12:12 h or shorter light periods.
One possible explanation of the retardation of growth and feed efficiency of tilapia fry
during shorter light periods in the present study is that during these short light phases
insufficient time was available for the establishment of a robust rhythmicity, as has been
reported by Biswas and Takeuchi (2002) and Biswas et al. (2002). The effect of photoperiod
on synchronizing an endogenous rhythm to the external environment may also require more
energy in the shorter light periods, leading to a reduction of somatic fish growth (Biswas
and Takeuchi, 2002; Biswas et al., 2002).

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399

The improvement in the performance of Nile tilapia fry in the present study with
increasing light period may also have been related to the reduction of standard metabolic
rate. In support, Biswas et al. (2002) and Biswas and Takeuchi (2002) studied the effects of
photoperiod on the metabolic rate of fed and unfed young and adult Nile tilapia. They found
that metabolic rate and energy loss were negatively correlated with light periods. They
concluded that Nile tilapia conserve energy when raised under photoperiods with longer
light phases. However, these authors suggested that growth studies must be conducted
under different photoperiod cycles in order to further evaluate the effects of photoperiod
regimes on these fish. The reduction of fish metabolic rate with increasing light phases has
also been reported with marine fish species (Boehlert, 1981).
The present study demonstrated that the growth and feed efficiency of fingerling Nile
tilapia were not significantly affected by photoperiod. The significant effect of photoperiod
on larval stages, but not fingerling stages, has also been reported with several other species,
including sole S. solea (Fuchs, 1978), black porgy M. macrocephalus (Kiyono and Hirano,
1981), yellow tail flounder Pleuronectes ferruguineus (Purchase et al., 2000) and
barramundi L. calcarifer (Barlow et al., 1995).
In contrast to the present results, Lourenco et al. (1998) reported that Nile tilapia
fingerlings maintained on 14 h light had better weight gain than those reared at 10 h light.
This controversy may have been related to the differences in culture systems, fish size, sex
ratio and light type and intensity. It is clear that more work is needed to verify the effects of
photoperiods on different sized-tilapia. The improvement in performance of fish juveniles
with extended light phases has also been documented with several other species, including
Atlantic salmon Salmo salar (Saunders et al., 1985), Atlantic cod Gadus morhua (Folkvord
and Ottera, 1993), turbot Scophthalmus maximus (Imsland et al., 1995, 1997) and haddock
Melanogrammus aeglefinus (Trippel and Neil, 2002).
Tilapia are generally diurnal feeders in nature and under culture conditions. They feed at
different times of the day, depending on species and size. Harbott (1975) found that Nile
tilapia in Lake Rudolf (Uganda) exhibit a regular diurnal feeding rhythm with maximum
feeding intensity between 5:00 and 8:00 a.m., and cease feeding between 14:00 and 18:00
p.m. Diurnal feeding activity has also been reported in cultured tilapia. Yousuf Haroon et al.
(1998) found that feeding activity of tilapia hybrids (O. mossambicus  O. niloticus) reared
in ponds was continuous during daytime, with a single feeding peak at around afternoondusk. The maximum feeding activity of O. niloticus  O. mossambicus hybrids reared in a
closed system occurred also during the afternoon Zavyalov and Lavrovskii (2001).
Feeding activity of Nile tilapia adults reared under conditions of self-feeding was also
observed exclusively during the light phases (Toguyeni et al., 1997). These studies clearly
pointed out the role of photoperiod on feeding activity and growth of these fishes.
Nile tilapia in the present study were fed the test diet three times per day, except for the
6L:18D group which was fed twice a day, due to the short light phase. During a 6-h light
period, feeding the fish three times (once every 2 h) may lead to feed waste, since the fish
may not accept the feed, as their stomachs will probably be still full from previous meals. In
addition, a number of studies indicated that feeding tilapia two to three times a day has
resulted in similar growth and FCR. Siraj et al. (1988) found that the best growth and FCR
of red tilapia (O. mossambicus  O. niloticus) fed at varying frequencies were attained at 2
and 3 feedings/day, and there was no significant difference between these two feeding

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A.-F.M. El-Sayed, M. Kawanna / Aquaculture 231 (2004) 393402

frequencies. Moreover, De Silva et al. (1986) found that the best performance of Nile tilapia
fingerlings fed a restricted ration at 3%/day was achieved at 1 or 2 feedings/day. Therefore,
one may argue that feeding frequency may have not significantly affected the present
results. Since fish fry and fingerlings were also given fixed rations (not ad libitum), at fixed
times, it is unlikely that feed consumption has been directly affected by photoperiod.
Changing metabolic hormone activities as a result of photoperiods may have been the prime
factor affecting fish performance, as suggested by Porter et al. (2001).
In conclusion, the present results revealed that photoperiods significantly affect the
growth of Nile tilapia fry, but not mixed-sex fingerlings. A 18L:6D cycle was suggested for
optimal performance of fish fry, while shorter light phases could be used in case of fish
fingerlings, taking into consideration the cost of electricity. These results have a significant
application in tilapia aquaculture in indoor recirculating systems, as they improve our
understanding of the role that photoperiod plays in fish growth and metabolism. Adopting
the optimum photoperiod in case of tilapia fry will also reduce the amount of energy used
for standard metabolism, and in turn increase fish growth and profitability.

Acknowledgements
The authors thank Dr. Salih A. Al-Shorapy, Associate Professor of Animal Genetics,
Department of Aridland Agriculture, College of Food Systems, United Arab Emirates
University, for running the statistical analyses of the results.

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