J. Appl. Ichthyol.

24 (2008), 31–37 Received: March 27, 2006

 2007 The Authors Accepted: December 10, 2006
Journal compilation  2007 Blackwell Verlag, Berlin doi: 10.1111/j.1439-0426.2007.00897.x
ISSN 0175–8659

Effect of feeding different protein to energy (P⁄ E) ratios on the growth performance
and body composition of Oreochromis niloticus fingerlings
By A. Ali1, S. M. Al-Ogaily2, N. A. Al-Asgah3, J. S. Goddard4 and S. I. Ahmed4
1
Department of Food Science and Nutrition, College of Agricultural and Marine Sciences, Sultan Qaboos University, Sultanate of
Oman; 2Natural Resources and Environmental Research Institute, King Abdulaziz City for Science and Technology, Riyadh, Saudi
Arabia; 3Zoology Department, College of Science, King Saud University, Riyadh, Saudi Arabia; 4Department of Marine Science
and Fisheries, College of Agricultural and Marine Sciences, Sultan Qaboos University, Al-Khod, Sultanate of Oman

Summary adequate supply of energy should therefore be provided from

This study evaluated the growth performance and body
composition of Oreochromis niloticus fingerlings (average
initial weight 16.53 ± 0.44 g) fed 9 experimental diets (A, B,
C, D, E, F, G, H and I) containing three different levels of
protein (26, 31 and 36 g 100 g)1) at three different gross energy
(GE) levels (16, 19 and 22 MJ kg)1) for a period of 64 days.
Significant differences were observed in the feed consumption,
body weight gain, specific growth rate (SGR), condition factor
(k), feed conversion ratio (FCR), protein efficiency ratio
(PER), net protein retention (NPR) and apparent net energy
retention (ANER) values of fish when the energy level of diet
was increased at different protein levels. The maximum weight
gain, SGR and k were observed on diet F containing 36%
protein and an energy level of 19 MJ kg)1 of dry feed with a
protein to energy (P ⁄ E) ratio of 18.96 (g protein MJ)1 GE).
A further increase in the energy content of the diet
(22 MJ kg)1) at the same protein level (Diet I) did not
produce any improvement in the growth performance. Low-
ering the energy level at the same protein level significantly
affected the growth performance. Fish fed diet B containing
31% protein and a lower energy level of 16 MJ kg)1 with the
same P ⁄ E ratio of 18.61 as diet F showed significantly lower
weight gain and growth performance than diet F. Diets E and
H containing 31% crude protein at all three energy levels
produced similar results as diet B. The poorest FCR was
observed when the diet contained both lower levels of protein
and energy. Fish fed diet G, containing 26% protein at high
energy level (22 MJ kg)1), showed the best PER and NPR
values. The PER and NPR were the poorest on diet C
containing 36% protein at low energy level (16 MJ kg)1). The
body moisture content at all protein levels decreased
(P < 0.05) with the increasing level of dietary energy whereas
the body fat content increased (P < 0.05). Similar trends were
observed in the body ash and energy content. Increasing the
dietary energy content at lower protein levels did not show any
difference (P > 0.05) in body protein content. Our results
indicated the optimum P ⁄ E ratio for O. niloticus as 18.96 g
protein per mega joule of gross energy at 36% dietary protein
level and a dietary gross energy value of 19 MJ kg)1.
Introduction
Fish in general eat to satisfy their energy demand. If the diet
does not contain sufficient energy, protein is used for energy
rather than for growth (Cowey and Sargent, 1979). An
non-protein energy sources. This can minimize the use of
costly protein and reducing the overall cost of fish production,
as protein is the single most expensive ingredient in finfish diets
(Watanabe, 2002; Ai et al., 2004). Each species of fish, even
those that are closely related, have slightly different dietary
requirements, at least for some essential nutrients. The optimal
protein requirements may vary with fish species, size, water
temperature and quality, variation in diet formulation as well
as culture system (Page and Andrew, 1973; NRC (National
Research Council), 1993; Al-Asgah and Ali, 1997; Al-Hafedth,
1999; Ai et al., 2004). The utilization of dietary protein is
related to both the protein level and the availability of non-
protein energy sources. The use of non-protein energy in fish
diets, however, must be closely evaluated as its excessive use
can reduce the feed intake, produce fatty fish and inhibit the
utilization of other nutrients (Ali and Al-Asgah, 2001; Hemre
et al., 2002). The optimization of protein to energy ratios
(P ⁄ E) in fish diets is widely accepted as a means to spare
protein, thereby increasing growth at a reduced cost (Lee and
Putnam, 1973; Machiels and Henken, 1985; Shiau and Peng,
1993; Lupatsch et al., 2001; Ali and Jauncey, 2005; Tibbetts
et al., 2005). An inadequate dietary P ⁄ E ratio may result in
lower growth as well as low protein and energy utilization
(Shiau and Lan, 1996; Samantaray and Mohanty, 1997;
Ai et al., 2004). It has been shown that at low P ⁄ E ratios,
the use of dietary protein for growth and maintenance of body
protein is maximized, while at higher P ⁄ E ratios, more protein
is used for energy or stored as fat. The protein sparing effect
occurs only if the minimum protein requirements are met,
including adequate amounts of amino acids (Van der Meer
et al., 1997).
A number of studies have been conducted to determine the
P ⁄ E ratio for some of the cultured fish species (Garling and
Wilson, 1976; Reis et al., 1989; Webster et al., 1995; Shiau and
Lan, 1996; Samantaray and Mohanty, 1997; Company et al.,
1999; Thoman et al., 1999; Hernandez et al., 2001; Ai et al.,
2004; Sahli et al., 2004; Ali and Jauncey, 2005). Mazid et al.
(1979) reported that the growth rate in young Tilapia zillii
increased proportionately to protein level in purified casein-
based diets up to an incorporation level of 35%. Any further
increase beyond 35%, however, gradually reduced the growth
rate. They concluded that diet containing about 30% crude
protein with a protein to energy (P ⁄ E) ratio of 81 mg
protein kcal)1 gross energy (GE) appeared to be more
efficiently utilized by fish in terms of protein deposition and

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32 A. Ali et al.

energy retention than diets with higher levels of crude protein.
El-Sayed and Teshima (1992) reported that at dietary protein
levels of 30–40%, fish (Oreochromis niloticus, fry) growth and
survival rates significantly improved with increasing dietary
energy levels. The best growth rate and feed utilization were,
however, achieved at 45% protein with a protein to energy
ratio of 110 mg protein kcal)1 GE. At the 50% dietary protein
level no further improvement in fish growth and feed
utilization were evident at any energy level. However, the
available information is too limited to establish the optimal
dietary protein to energy (P ⁄ E) ratio in the diets of O. niloticus
fingerlings. The present study was therefore conducted to
determine the optimum protein, energy and protein to energy
ratio in practical diets for O. niloticus fingerlings.
Materials and methods
Oreochromis niloticus fingerlings (mixed sex) with a mean
weight of 16.53 ± 0.44 g were obtained from the fish
hatchery of King Abdulaziz City for Science and Technol-
ogy (KACST) Deerab, Riyadh, Saudi Arabia. The fish were
acclimatized to the experimental conditions for a period of
2 weeks before the start of the actual experiment. During
this period they were fed the same standard diet (Fish Feed-
Larvae Code #619, Grain Silos Organization, Riyadh) as
used previously at the hatchery. At the start of the
experiment thirty randomly captured fish (divided into three
replicates of 10 fish in each) were killed rapidly by
immersion in ice water. Body weight and length were
determined prior to storage at )30C for determination of
initial body composition at a later time. Two hundred
seventy fish were randomly divided into 27 replicates with 10
fish in each. Nine different experimental diets (A, B, C, D,
E, F, G, H and I) were then randomly assigned to 3
replicates containing 10 fish in each replicate. These
comprised 9 treatment groups (A–I), each with 3 replicates
of 10 fish per aquarium. The fish were kept in glass tanks
(100 · 42.5 · 50.0 cm) containing dechlorinated, well-aerated
tap water and fitted with a waste filtration facility. The
water temperature was maintained at 28 ± 1C with the
help of a thermostatically controlled heating system.
Compressed air was used to maintain the oxygen supply.
Water quality parameters were monitored every 2 weeks.
The values recorded were: pH (7.1–8.0), dissolved oxygen
(5.6–6.7 mg L)1), ammonia nitrogen (0.12–0.20 mg L)1),
nitrite nitrogen (0.33–0.58 mg L)1) and alkalinity as CaCO3
(235–350 mg L)1).
Nine experimental diets containing 3 different levels (26%,
31% and 36%) of crude protein (CP) at 3 different gross
energy (GE) levels (16, 19 and 22 MJ kg)1) with different
protein to energy (P ⁄ E) ratios (ranging from 12 to 22 g
protein MJ)1 GE) were prepared using a pellet press with
2 mm die (Table 1). The diets were dried at 60C and stored at
)18C throughout the experimental period. The proximate
chemical composition of the experimental diets was deter-
mined by the methods of AOAC (Association of Official
Analytical Chemists) (1995) and is given in Table 2. Gross
energy content of the diets was calculated on the basis of their
crude protein, total fat (ether extract) and carbohydrate
(nitrogen free extract, NFE) contents using the equivalents
of 23.64, 39.54, and 17.15 MJ kg)1, respectively (Kleiber,
1975). Each diet was allotted to 3 replicates in a completely
randomized design. The diets were offered ad libitum twice
daily to satiation for a period of 64 days. Daily feed intake and
fortnightly weight gains were recorded. To quantify the exact
amount of dry feed intake, any feed refused was immediately
siphoned out, dried and weighed. The experiment was
conducted under artificial light with a light : dark cycle of
12 : 12 h. At the end of the experiment all fish were killed and
their body weight and length recorded. To determine the whole
body composition, fish were cut into pieces, minced in a meat
grinder, homogenized and immediately frozen at )30C for
further analysis. The proximate chemical composition was
determined according to the methods of AOAC (Association
of Official Analytical Chemists) (1995). The gross energy (GE)
content of fish was calculated from the total fat and crude
protein contents using the equivalents of 39.54 MJ kg)1 crude
fat and 23.64 MJ kg)1 crude protein (Kleiber, 1975). Feed
conversion ratio (FCR), specific growth rate (SGR), protein
efficiency ratio (PER), net protein retention (NPR), and
apparent net energy retention (ANER) values were calculated
as follows:

Table 1
Composition of experimental diets (%)

Experimental diets

Low energy Medium energy High energy

Ingredients A B C D E F G H I

Fishmeal 18.00 25.00 30.00 18.00 25.00 30.00 19.00 27.00 32.00
Soybean meal 17.00 20.00 25.00 17.00 19.00 25.00 18.00 19.00 25.00
Maize 41.00 31.00 21.00 43.00 34.00 23.00 27.00 18.00 7.00
Wheat bran 12.00 12.00 12.00 12.00 12.00 12.00 12.00 12.00 12.00
Corn oil – – – 3.00 3.00 3.00 10.00 10.00 10.00
Cod-liver oil – – – 3.00 3.00 3.00 10.00 10.00 10.00
Gelatin 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00
Mineral mixture1 2.00 2.00 2.00 2.00 2.00 2.00 2.00 2.00 2.00
Vitamin mixture2 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00
a-Cellulose 8.00 8.00 8.00 – – – – – –
Total 100.00 100.00 100.00 100.00 100.00 100.00 100.0 100.00 100.00
1
Per kg of premix contains: CaHPO4, 530 g; K2HPO4, 80 g; Na2HPO4, 90 g; MgCl2Æ6H2O, 100 g; KCl, 67.5 g; K2SO4, 80 g; NaCl, 30 g;
KI, 0.05 g; ZnSO4Æ7H2O, 2.5 g; SeO2, 0.03 g; CuSO4Æ5H2O, 0.15 g; FeSO4Æ7H2O, 18 g; (NH4)6Mo7O24Æ4H2O, 0.01 g; MnSO4ÆH2O, 0.5 g; NaF,
1.2 g; CoCl2Æ6H2O, 0.01 g.
2
Per kg of premix contains: vitamin A, 400 000 IU; D3, 200 000 IU; E, 5000 IU; K3, 1 g; B1, 1 g; B2, 1.5 g; B6, 1 g; pantothenic acid, 5 g; niacin,
3 g; folic acid, 0.5 g; B12, 2 mg; biotin, 100 mg; vitamin C, 20 g.
Effect of feeding different protein to energy (P ⁄ E) ratios 33

Table 2
Proximate chemical composition of the experimental diets (on % dry matter basis)

Experimental diets

Low energy Medium energy High energy

Parameters A B C D E F G H I

Dry matter (%) 93.36 93.81 93.93 93.26 93.09 93.59 94.14 94.66 94.18
Crude protein 26.40 30.88 36.59 26.33 31.46 36.76 25.92 30.82 36.38
Crude fibre 10.47 10.61 10.39 3.59 3.38 3.51 3.71 3.63 3.85
Total fat 3.64 3.72 4.12 9.36 9.69 9.64 24.20 24.15 24.04
Ash 8.02 9.04 10.07 8.18 9.34 10.17 8.07 9.13 10.35
Nitrogen-free extract 51.47 45.75 38.83 52.54 46.13 39.92 38.10 32.27 25.38
Gross energy (MJ kg)1) 16.50 16.59 16.97 18.99 19.16 19.39 22.17 22.35 22.45
Gross energy (kcal kg)1) 3929 3950 4040 4521 4562 4617 5279 5321 5345
P ⁄ E (g protein MJ)1GE) 16.00 18.61 21.56 13.87 16.42 18.96 11.69 13.79 16.20
P ⁄ E (mg protein kcal)1GE) 67 78 92 60 69 82 48 58 68

Feed conversion ratio (FCR) (1989). To determine the growth rate as a function of dietary
¼ kg dry feed consumed per kg wet weight gain: protein content, where optimal, with a corresponding dietary
energy value, the growth data was analyzed by stepwise
9 regression analysis using the response surface. The regression
Specific growth rate (SGR), >
= 100 [ln final wt (g) analysis was done using MINITAB PROGRAM V.13 (Minitab Inc,
as percentage of body weight ¼  ln initial weight (g)]
>
; 2003).
gain per day =time (days):

Protein efficiency ratio (PER) Results

¼ live weight gain (g)/protein consumed (g)
Net protein retention (NPR)
¼ [increase in carcass protein/protein fed]  100
Apparent net energy retention (ANER)
¼ [(final body GE KJ g1 Þ
 ðinitial body GE KJ g1 Þ=KJ GE fed  100
The body condition factor (k) was calculated according to the
equation k = [W(g) ⁄ L(cm)3] · 100, where W is the wet
weight of fish in grams and L is the length in centimeters.
The results were subjected to statistical analysis using the
analysis of variance technique and the means were compared
by Fishers LSD test according to Snedecor and Cochran
The data on the growth performance of O. niloticus fingerlings
fed 3 different levels of protein (26%, 31% and 36%) at 3
energy levels (16, 19 and 22 MJ kg)1 GE) constituting different
protein to energy ratio is presented in Table 3. No fish
mortality was observed in any of the groups during the entire
experimental period. Significant differences were observed in
the body weight gain, SGR and k of fish when the energy level
of diet was increased at different protein levels. The maximum
weight gain, SGR and best k were observed on diet F
containing 36% protein at an energy level of 19 MJ kg)1 of
dry feed with a protein to energy (P ⁄ E) ratio of 18.96
(g protein MJ)1 GE). A further increase in the energy content
of the diet (22 MJ kg)1) at the same protein level (Diet I) did
not produce any improvement in the growth performance of
fish. Lowering the energy level at the same protein level,
however, significantly affected the growth performance of fish.

Table 3
Growth performance of Oreochromis niloticus fed different levels of protein to energy (P ⁄ E) ratios

Experimental diets

Low energy Medium energy High energy

Parameters A B C D E F G H I SE

Initial weight (g fish)1) 16.75 16.67 16.23 16.65 16.71 16.31 16.35 16.32 16.82 0.44NS
Final weight (g fish)1) 49.00g 54.25d 54.17d 51.16f 57.31c 61.69a 52.88e 53.36de 60.15b 1.21
Weight gain (g fish)1) 32.25g 37.58de 37.94d 34.51f 40.60c 45.38a 36.53e 37.04de 43.33b 1.26
Specific growth rate (SGR) 1.69d 1.85bc 1.91b 1.76dc 1.92b 2.08a 1.84bc 1.85bc 2.00ab 0.09
Condition factor (k) 2.65c 2.91b 2.84bc 2.71c 3.02ab 3.18a 2.87bc 2.93b 3.11a 0.18
Feed consumed (DM g fish)1) 47.10b 48.23a 48.34a 48.46a 48.12a 47.52ab 48.60a 46.44b 46.94b 1.13
Feed conversion ratio (FCR) 1.46a 1.28bc 1.27bc 1.40ab 1.18bc 1.05d 1.33b 1.25bc 1.08d 0.12
Protein efficiency ratio (PER) 2.54c 2.52c 2.14d 2.70b 2.68b 2.60bc 2.89a 2.59bc 2.54c 0.14
Net protein retention (NPR) 37.39e 38.70cd 32.87e 40.16bc 41.03ab 40.23bc 41.43a 39.13c 37.25d 1.67
Apparent net energy retention 18.14c 24.93bc 24.59bc 21.14d 25.24b 27.29a 23.67c 21.82d 24.74bc 1.53

SE, pooled standard error; NS, non significant.

a–g
Different superscripts in same row = significant differences between values at 5% (n = 3).
34 A. Ali et al.

Table 4
Proximate body composition of Oreochromis niloticus fed different levels of protein to energy (P ⁄ E) ratios (on % wet basis)1

Experimental diets

Low energy Medium energy High energy

Parameters A B C D E F G H I SE
a a a b c bc c d cd
Moisture 75.32 75.14 75.77 74.10 73.13 73.79 73.01 72.23 72.60 1.07
Crude protein 14.52bc 15.08a 14.98ab 14.74bc 15.07ab 15.14a 14.67c 14.83b 14.54c 0.31
Total fat 3.95e 3.89e 3.98e 4.62c 4.67c 4.38d 5.08b 5.47a 5.57a 0.14
Ash 4.33d 4.64b 4.69b 4.48c 4.73b 4.46c 4.99a 4.76b 5.03a 0.11
Gross energy (MJ kg)1) 5.00d 5.10d 5.11d 5.31bc 5.41bc 5.30c 5.47b 5.66a 5.63a 0.15
1
Composition of fish slaughtered at the beginning of the experiment (moisture, 77.02%; crude protein, 14.17%; fat, 3.23%; ash, 4.95%; and gross
energy, 4.63 MJ kg)1).
SE, pooled standard error; NS, not significant.
a–e
Different superscripts in same row = significant differences between values at 5% (n = 3).

The fish fed diet B containing 31% protein at a lower energy
level of 16 MJ kg)1 but having the same P ⁄ E ratio of 18.61 as
diet F showed significantly lower weight gain and growth
performance as compared to diet F. The diets E and H
containing 31% of crude protein at both medium (19 MJ kg)1)
and high energy level (22 MJ kg)1) produced similar results as
in diet B.
Increasing the energy level of diets at a lower protein level
(26%) showed significant improvement in the growth
performance. The fish fed diet F showed the best FCR
value (1.05). The poorest FCR was observed at both lower
dietary protein and energy levels. Increasing the protein level
(26–36%) at the same energy level, however, significantly
improved the efficiency of feed conversion ratio. Increasing
the dietary protein content with increasing level of dietary
energy showed significant improvement in the FCR; how-
ever, the differences between the medium and high energy
levels at the highest dietary protein level were non-
significant. At the same dietary energy level, the PER
proportionately decreased with the increase in dietary
protein content. The best PER value was observed at 26%
dietary protein at an energy level of 22 MJ kg)1. Increasing
the dietary energy level at lower dietary protein levels
showed better PER values as compared to higher dietary
protein levels. No significant (P < 0.05) differences were
observed in PER values at 31% and 36% dietary protein
levels at medium and high energy levels of the diet. Similar
results were observed with the NPR values. The ANER
values were the best in diet F containing 36% dietary
protein at an energy level of 22 MJ kg)1 with a protein to
energy ratio of 18.96. Increasing the energy level from low
to medium at all dietary protein levels increased the ANER
values. A further increase in the energy level (22 MJ kg)1) of
diets at all 3 dietary protein levels did not (P > 0.05)
increase the ANER values.
Significant (P < 0.05) differences were observed in the
proximate body composition of the O. niloticus fed different
levels of protein to energy ratios. The body moisture content at
all protein levels decreased significantly (P < 0.05) with the
increasing level of dietary energy (Table 4). Increasing the
dietary energy content at the lower protein level did not show
any significant difference (P > 0.05) in body protein content,
whereas at all protein levels the increasing dietary energy level
significantly (P < 0.05) increased body fat content. Similar
trends were observed in body ash and energy content of fish.
To determine the growth rate as a function of dietary
protein content, where optimal, with a corresponding dietary
energy value, the growth data was analyzed by stepwise
regression analysis using the response surface. The growth and
nutrient utilization data indicated that increasing the level of
dietary protein produced significantly (P < 0.05) greater
responses than the dietary energy levels. The following
regression equations were drawn:
Specific Growth Rate (SGR) ¼ 0:909 þ 0:0222CP
þ 0:0142GE ðr2 ¼ 82:6%Þ
FeedConversion Ratio (FCR) ¼ 2:43  0:0248CP
 0:0205GE ðr2 ¼ 83:2%Þ
Protein Efficiency Ratio (PER) ¼ 2:61  0:0285CP
þ 0:0445GE ðr2 ¼ 67:8%Þ
Discussion
The main objective of this study was to determine the optimum
dietary P ⁄ E ratio and to establish at which dietary protein and
energy levels the growth performance of fish is maximized. The
optimization of protein to energy ratios (P ⁄ E) is widely
accepted as a means to spare protein, thereby increasing
growth at a reduced cost (Lee and Putnam, 1973; Machiels and
Henken, 1985; Shiau and Peng, 1993; Company et al., 1999;
Ai et al., 2004; Sahli et al., 2004). The maximum weight gain,
SGR, and best FCR as well as k were observed in diet F
containing 36% protein at a medium energy level of
19 MJ kg)1 of dry feed with a protein to energy (P ⁄ E) ratio
of 18.96 (g protein MJ)1 GE). Lowering the dietary crude
protein contents both at lower or higher dietary energy levels
but having the same dietary P ⁄ E ratio decreased the growth
performance of fish. The results of the present study indicated
that even at the same P ⁄ E ratio, if the required protein and
energy levels are not maintained, then the protein may be
utilized for energy rather than for growth. Reducing the P ⁄ E
ratio with energy dense diets did not improve the efficiency of
feed utilization or growth performance of fish. An optimum
P ⁄ E ratio can be obtained at lower dietary protein and energy
levels, however it will not be effective in obtaining the
maximum growth performance until the minimum dietary
protein and energy requirements of the fish are met. Any
inadequate dietary P ⁄ E ratio may result in lower growth as
well as low protein and energy utilization (Shiau and Lan,
1996; Samantaray and Mohanty, 1997; Ai et al., 2004). It has
been shown at low P ⁄ E ratios that the use of dietary protein
Effect of feeding different protein to energy (P ⁄ E) ratios
for growth and maintenance of body protein is maximized,
while at higher P ⁄ E ratios more protein is used for energy or
stored as fat. The protein sparing effect occurs only if the
minimum dietary protein requirements are met, including
adequate amounts of amino acids and extra dietary protein
used for energy purposes (Van der Meer et al., 1997). The
relatively higher percentage of dietary crude fibre content at
lower dietary energy levels may also be regarded as one of the
factors for the poor performance of fish as it may interfere with
the digestion of other nutrients (Al-Asgah and Ali, 1996;
Alexis, 1997). Lipid is regarded as a highly available source of
energy (NRC (National Research Council), 1993), however the
digestible energy content of carbohydrates may vary among
fish species (Krogdahl et al., 2005). The non-protein energy
source can either be carbohydrate or lipid and an optimum
carbohydrate to lipid ratio in the diets of O. niloticus has
already been established (Ali and Al-Asgah, 2001). Mazid
et al. (1979) observed that diet containing about 30% of crude
protein with a protein to energy ratio (P ⁄ E) of 81 mg
protein kcal)1 GE appeared to be more efficiently utilized by
the fish (T. zillii) in terms of protein deposition and energy
retention than diets with higher levels of crude protein.
El-Sayed and Teshima (1992) reported that the best growth
rate and feed utilization in Nile tilapia (O. niloticus) fry were
achieved at 45% protein level at 400 kcal GE 100 g)1 diet with
a protein to energy ratio of 110 mg protein kcal)1 GE.
Kaushik et al. (1995) examined the protein and energy needs
of juvenile O. niloticus. They reported that the protein intake
for the maximum carcass nitrogen gain was 12 g kg)1 and the
maintenance level was 2 g kg)1 day)1 at an optimal protein to
energy ratio of 18 mg kJ)1. Our results are in line with these
findings and also with the findings of Teshima et al. (1985)
who concluded that Tilapia nilotica fingerlings grew optimally
on diets containing 30–40% of protein at energy levels of 17 to
22 MJ kg)1. El-Saidy and Gaber (2005), however, reported no
significant increase in the growth rate of adult Nile tilapia
(O. niloticus) with increasing dietary protein levels (25 or 30%).
This may be due to a number of factors as each species of
fish, even those that are closely related, have slightly different
dietary requirements, at least for some essential nutrients. The
optimal value may vary with fish species, size, variation in diet
formulation and culture system (Ai et al., 2004).
The PER decreased when the dietary protein content was
increased at the same energy level. However, increasing the
dietary energy level at lower dietary protein levels improved
the efficiency of protein utilization and retention, indicating
that when adequate amounts of non-protein energy source
were available, the dietary protein was most efficiently used for
tissue growth and body protein synthesis. Increasing the
energy level from low to medium at all dietary protein levels
increased the apparent net energy retention values. Overall, the
increase in dietary crude protein content (26–36%) signifi-
cantly improved the performance of fish but the simultaneous
increase in dietary energy levels (16–22 MJ kg)1) only
produced significant improvement up to medium energy level
(19 MJ kg)1). This indicated that in energy dense diets, the
dietary protein content should only be increased at medium
dietary energy level to obtain an optimum P ⁄ E ratio for
increased performance. The dietary protein requirements must
be met if the non-protein energy source is to be optimally
utilized, otherwise the protein will be used for energy purposes.
Above the optimum protein requirements, high protein to
energy ratios result in a higher proportion of protein being
deaminated and used for energy needs or stored as fat.
35
A reduction in the digestible protein to energy ratio resulted in
decreased growth rates (Van der Meer et al., 1997). Protein
retention is influenced by a number of factors including
digestibility of feed ingredients, protein content of the diet,
amino acid balance and dietary protein to energy ratio. The
precise mechanism involved in the regulation of nutrient
partitioning and energy utilization needs to be fully estab-
lished, however the use of high-energy diets, distributed in
excess could compromise the growth rate and feed efficiency by
metabolic endocrine adjustments. Mathis et al. (2003) reported
that high-energy feed led to a reduction in the excretion of
nitrogen compounds in Eurasian perch (Perca fluviatilus).
Company et al. (1999) suggested that as far as P ⁄ E ratio is
concerned, the conclusions and recommendations should be
feeding rate dependent. Dietary protein constitutes one of the
primary nutrients that not only affects the cost of feed but also
is the initial source of nitrogen waste products entering the
culture system. Consequently, optimization of dietary protein
levels along with increasing nutrient retention by fish could
reduce the nitrogen loading and can positively influence the
production costs. Further studies are therefore required and
should be based on higher protein levels to determine at what
protein level the amino acid requirements are best met and
what is the optimum P ⁄ E ratio at these higher dietary protein
and energy levels.
Significant (P < 0.05) differences were observed in the
proximate body composition of the O. niloticus fed different
protein to energy ratios. The changes in carcass composition
were, however, less consistent. The body moisture content at
all protein levels decreased significantly (P < 0.05) with
increasing level of dietary energy. Increasing the dietary energy
content at a lower protein level, however, did not show any
significant differences (P > 0.05) in body protein content
whereas at all protein levels, increasing the dietary energy level
significantly (P < 0.05) increased the body fat content.
Similarly, with the increase in both dietary protein and energy
levels, a significant increase in the energy gains of fish was
observed. These results are in line with earlier findings of
El-Sayed and Teshima (1992). Kim and Lall (2001) reported
that the nitrogen intake by juvenile haddock (Melanogrammus
aeglefinus) increased with an increase in dietary protein
(45–65%), which in turn resulted in an increase in nitrogen
gain of fish whole body. Our results however are not in line
with findings of El-Saidy and Gaber (2005) who reported that
the dietary protein level (25–30%) did not affect the whole
body fat, protein, ash or energy content of fish. This may be
due to a number of factors including the diet formulation and
ingredient composition. Both the endogenous and exogenous
factors operate simultaneously to influence the body composi-
tion of fish (Haard, 1992; Shearer, 1994). Although the
available dietary energy level plays an important role in
determining body lipid composition, the dietary lipid content
is regarded as the most important factor influencing the
carcass lipid in fish (Sargent et al., 1989; Hanley, 1991; Ali and
Al-Asgah, 2001). Low protein ⁄ energy ratio is suspected as
affecting fat accumulation in the liver (Brown et al., 1996;
Ibeas et al., 1996). Mathis et al. (2003) did not observe any
relationship between the biochemical and organoleptic char-
acteristics of fish flesh and dietary treatment in growing
Eurasian perch (P. fluviatilis) of the same size and growth rate.
The results of the present study indicated that the optimum
P ⁄ E ratio for O. niloticus is 18.96 g protein MJ)1 of gross
energy at 36% dietary protein level with a dietary gross energy
value of 19 MJ kg)1.
36
Acknowledgements
We express our thanks to King Abdulaziz City for Science and
Technology (KACST), Riyadh, Saudi Arabia for providing
the fish and other facilities during this study. Thanks are also
due to Grain Silos and Floor Mills Organization, Riyadh, for
the supply of feed ingredients. We also acknowledge the help
provided by Mr. M. Sarwar Khan, Department of Statistics,
KSU, Riyadh and Dr M. Idrees Ahmad, Department of
Mathematics and Statistics, SQU, Muscat, Oman, during the
statistical analysis of the data. We also thank Dr Santosh
P. Lall, National Research Council, Institute of Marine
Biosciences, Halifax, Canada for reviewing the manuscript
and giving valuable suggestions for its improvement.
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Authors address: Dr Amanat Ali, Department of Food Science and
Nutrition, College of Agricultural and Marine
Sciences, Sultan Qaboos University, PO Box 34,
Al-Khod 123, Sultanate of Oman.
E-mail: amanat@squ.edu.om