African Journal of Pure and Applied Chemistry Vol. 5(6), pp.

127-135, June 2011

Available online at http://www.academicjournals.org/AJPAC
ISSN 1996 - 0840 ©2011 Academic Journals

Full Length Research Paper

Fatty acid profiles of the eggs and juvenile muscle of

Nile perch (Lates niloticus, L. 1758) caught from Lake
Victoria, Uganda
V. T. Namulawa1, Jolocam Mbabazi2* and J. Kwetegyeka3
1
Aquaculture Research and Development Centre, Kajjansi, P.O. Box 530, Kampala, Uganda.
2
Department of Chemistry, Makerere University, P.O. Box 7062, Kampala, Uganda.
3
Department of Chemistry, Kyambogo University, P.O. Box 1, Kyambogo, Uganda.
Accepted 5 April, 2011

Fatty acid profiling of the eggs and muscle of L. niloticus juveniles was carried out, revealing the
presence of two types of cholesterol and over twenty fatty acids. The total saturated fatty acid, total
monounsaturated acid and the total polyunsaturated fatty acid compositions in the various samples lay
in the ranges 13.39 to 41.81, 19.41 to 42.98 and 22.17 to 43.63%, respectively. There was a detection of
an abundance of palmitic, stearic, oleic and docosahexaenoic fatty acids in all the samples
investigated. The n-3/n-6 ratio averaged at 5.12±4.28. Cholesterol was detected in all the samples except
in the eggs. This information is necessary as an aid to the formulation of an entire feed process
required for the rapid replenishing of the declining L. niloticus stocks in Lake Victoria.

Key words: Lates niloticus, muscle tissue, fatty acids, Lake Victoria, cholesterol.

INTRODUCTION

Nile perch (Lates niloticus) are commonly found in the the Nile perch is largely fatty, so much so that local
waters of the River Nile and in the lakes and other rivers fishermen used to fry it all by itself and collect its oil for
in eastern, central and western Africa (Hopson, 1972). further domestic use. L. niloticus is known to be rich in
The fish species gained its popularity as a source of highly unsaturated and polyunsaturated fatty acids
protein in East Africa in the late 1970’s, some twenty (Ogwok et al., 2008), which renders the species an
years after its first introduction in the waters of the Lakes important source of these fatty acids. Fatty acid profile
Victoria, Nabugabo and Kyoga (Ogutu-Ohwayo, 1985; data in other fishes, plants and animal tissues is
Bwanika et al., 2006). Currently, Nile perch is a highly mentioned elsewhere in current literature (Turon et al.,
demanded East African fish export commodity (Okedi, 2005; Ugoala et al., 2009; Tang et al., 2009). Research
2005), exerting undue pressure on the fisheries industry findings indicate that one of the avenues through which
and resulting in a marked decrease in Nile perch Nile perch could possibly be sustainably conserved is
populations in the respective water bodies (LVFO, 2009). through aquaculture (Gregory, 2006), and that this could
The rather rapid decline in Nile perch populations in East go well alongside developing an artificial fish feed for this
Africa and in the waters of Lake Victoria in particular over species. In this respect, studies indicate a close
the recent past, has precipitated major concerns relationship between body composition and the dietary
regarding the urgent need to put in place technologies fatty acid requirements in fish (Mourente and Tocher,
geared towards the restoration of the Nile perch 1992a).
populations (Munyaho, 2004). Unlike many of the other Fish require fat as a source of energy and also as a
fish species commonly found in the Great Lakes region, source of fatty acids that are important ingredients in
growth and reproduction (Sargent et al., 1995). The level
of fat within the body of a fish highly depends on its
dietary levels as well as on the different stages in the life-
*Corresponding author. E-mail: cycle of the fish (Uotani et al., 1990). Fatty acids usually
[email protected]. comprise a chain of carbon atoms with a carboxyl group
128 Afr. J. Pure Appl. Chem.

at one end and a methyl group at the other end. In methanol. The gas was turned off when the methanol reached 2M
saturated fatty acids, each carbon atom in the molecular in HCl. Sub-samples of the fish eggs, egg-fat and muscle tissue
(weighing approximately 30 to 50 mg) were transferred to thick-
chain forms single bonds with four other atoms. If a
walled glass tubes. 1 ml of the acidified anhydrous methanol was
carbon is not bound to four other atoms, double or triple added. Nitrogen gas was flushed through the tubes which were
bonds are formed. Consequently, fatty acids are normally then securely sealed with Teflon-lined screw caps and left for a
categorised based on molecular chain length, the period of 2 h in an oven thermostated at 90°C. The samples were
existence of multiple bonds and the spatial arrangement subsequently methanolysed and all the fatty acids in the samples
of hydrogen atoms in these bonds (Abbas et al., 2009). were converted to their methyl esters in the methanolic solution
(Grahl-Nielsen and Barnung, 1985).
Fatty acids are important for fish growth, reproduction
and movement, as well as being a major source of
energy (Tocher, 2003). Fish adopt various nutritional Extraction of the fatty acid methyl esters
requirements at their different stages of development
(Tang et al., 2009). It has, for example, been observed in After cooling to room temperature, half the methanol was allowed to
L. niloticus (Hopson, 1972) that 0.30 to 0.35 cm larvae evaporate by bubbling nitrogen gas, with 0.5 ml distilled water being
added to reduce the solubility of the esters. Extraction of the
tend to feed on Cladocera and Copepod nauplii, whereas product from the methanol-water phase was effected using two
those between 0.35 and 1.00 cm feed primarily on successive 1.0 ml aliquots of HPLC grade n-hexane and
Copepods and insect nymphs. Consequently, during the centrifuged at 1500 rpm for 3 min. The methyl esters were obtained
culturing of L. niloticus it becomes of vital importance to from the upper organic layer by siphoning and stored under
provide feeds with the appropriate fatty acid inclusions at refrigeration for subsequent gas chromatograph/mass spectrometry
(GC/MS) analysis using an Agilent 6890N GC–MS (USA version)
the larval (Tang et al., 2009), juvenile (Mourente, 2003) spectrometer.
and broodstock stages (Sorbera et al., 2001), so as to
offset possible nutritional deficiencies and to increase the
chances of larval survival (Rimmer and Reed, 1989). GC/MS analysis of the methyl fatty acid esters
The purpose of the present study therefore was to
establish the fatty acid profiles of the eggs as well as of 1 µl of the mixed hexane extracts was injected onto a 25 by 0.25
mm fused silica column with polyethylene-glycol (PEG) as the
the muscles of L. niloticus at various stages of growth. stationary phase with a 0.2 µm thickness (CP-WAX 52CB
This knowledge would subsequently be used to provide Chrompack) and helium gas at 20 psi as the mobile phase. The
vital information on the fatty acid dietary requirements for column was mounted in a GC/MS (Agilent 6890N). The column was
L. niloticus at its various stages of development, and as mounted in a GC/MS (Agilent 6890N with autosampler 7683B
an aid in the feed formulation process. This may be series, fitted with an electronic pressure control and mass selective
important during the culture of L. niloticus, which is an detection (ionisating energy, 70 eV; source temperature, 250°C)).
The injector temperature was 260°C. The temperature of the
avenue for the rapid replenishing of L. niloticus stocks in column was kept at 90°C for 4 min after injection and thereafter
Lake Victoria. increased to 165°C at a rate of 30°C/min, followed by an increase
of 3°C/min to 225°C. The temperature was then maintained at
225°C for close to 11 min. The fatty acids in the samples were
MATERIALS AND METHODS identified by means of the standard mixture GLC-68D from Nu-
Chek-Prep (Elysian, Minn., USA) containing 20 fatty acids and by
Sample collection and analysis mass spectrometry. Quantification of the esters was achieved by
integration of the peaks using Chemstation software obtained from
Twelve fish of various sizes were caught from Lake Victoria at Thermo LabSystems, with the relative amount of each fatty acid
Kiggungu landing site (Longitude 36° 26′ 15″E, Latitude 00° 02′ ester in each sample being expressed as a percentage of all the
49″N) using 3" beach seine and 7 mm mosquito nets. The samples esters in the sample.
were divided into four groups according to the total length: 0 to 10,
11 to 20, 21 to 30 and 31 to 40 cm, with each group comprising
three members. The fish were immediately transported in ice- Statistical analysis
cooled boxes to the Department of Chemistry, Makerere University,
for laboratory analysis. The eggs were collected from three sexually The results obtained from the chromatography readings were
mature females (92.5 to 97.0 cm total length). Muscular portions presented as means ±SD. The data was analysed using one-way
from the caudal and abdominal areas were sliced off the fish using ANOVA and the correlations were performed using
a sharp clean stainless-steel knife. The eggs, egg fat and muscle GraphPad.Prism.v5.01 Statistical software.
were separately homogenised using a blender and stored at −86°C
for 12 h, before further analysis. In each case, the analyses were
carried out in triplicate. RESULTS

Esterification of the fatty acids A total of twenty-seven different fatty acids along with two
cholesterol isomers were detected from the eggs, egg fat
Dry hydrogen chloride gas was bubbled through anhydrous and muscle tissue (Tables 1, 2 and 3). The total
methanol (HPLC grade) in a flask immersed in an ice-bath and its saturated fatty acid composition ranged from 13.39±2.45
concentration periodically monitored by the increase in mass of the to 40.81±0.46%, that of the monounsaturated acids from
 Namulawa et al. 129

Table 1. Fatty acid content (expressed as percentage of total fatty acids) in the eggs and fat tissue
around the eggs of L. niloticus.

F/A mean percentages

Fatty acids Egg fat N = 3 Egg N = 3
12:0 0.16±0.00 0.02±0.00
14:0 4.45±0.26 1.20±0.25
15:0 0.49±0.00 0.10±0.03
16:0 23.43±0.54 8.37±1.60
17:0 1.49±0.04 0.31±0.05
18:0 10.16±0.30 3.39±0.51
Total saturated fatty acids 40.18±0.46 13.39±2.45
14:In5 0.13±0.00 0.05±0.00
iso 15:00 0.68±0.00 0.27±0.06
16:In7 7.34±0.21 6.26±1.22
iso 17:00 0.72±0.05 0.47±0.01
α 1so 17:00 0.26±0.03 0.11±0.02
17:1n9 1.09±0.04 0.93±0.14
18:1n9 21.82±0.53 31.12±9.28
18:1n7 4.96±0.11 3.53±1.13
20:1n9 0.64±0.02 0.24±0.04
Total monounsaturated fatty acids 37.64±0.35 42.98±11.87
18:2n6 4.76±0.26 2.15±3.04
18:2n4 0.54±0.04 14.05±19.21
18:3n6 0.22±0.01 2.07±2.09
18:3n3 4.92±0.18 7.18±1.09
20:3n6 0.53±0.04 1.02±0.15
20:4n6 1.64±0.06 2.66±0.40
20:5n3 (EPA) 1.07±0.05 1.04±0.16
21:5n3 0.23±0.01 0.00±0.00
22.4n6 0.59±0.03 0.48±0.00
22:5n6 0.49±0.01 0.95±0.13
22:5n3 3.04±0.07 3.09±0.44
22:6n3 (DHA) 4.13±0.07 8.95±1.57
DHA/EPA 3.86 7.38
Total polyunsaturated fatty acids 22.16±0.81 43.64±14.32
C1 N.D N.D
C2 N.D N.D
Total cholesterol 0.00 0.00
(n - 3) total 13.39 20.26
(n - 6) total 8.23 9.33
n-3/n-6 1.63 2.17
N.D: Not detected.

18.73±0.25 to 42.98±11.87%, while that of the acids, it was oleic acid (18:1n9) that was the most
polyunsaturated fatty acids lay between 22.16±0.81 and abundant (Tables 2 and 3). Of the polyunsaturated fatty
43.64±14.32%. Of the saturated acids, the most acids, docosahexaenoic (C22:6n3) was the most
abundant was palmitic acid (C16:0) occurring in the egg available. The analysis also indicated that 16:0, 18:0,
fat, 21 to 30 cm of the abdominal muscle and 31 to 40 cm 16:1n7, 18:1n9, 18:1n7, 18:3n3, 18:2n6, 20:4n6, 20:5n3,
of the caudal muscle (23.43±0.54, 24.53±0.01 and 22:5n3 and 22:6n3 (Figure 1) were the most abundant
23.75±0.86%, respectively). Among the monounsaturated fatty acids in the eggs and all the other portions of the
130 Afr. J. Pure Appl. Chem.

Table 2. Fatty acid content (expressed as percentage of total fatty acids) in the abdominal muscle of L. niloticus.

Fish size classes (cm)

Abdominal
0-10 11-20 21-30 31-40
Fatty acids N=3 N=3 N=3 N=3
12:0 0.15±0.04 0.18±0.02 0.13±0.00 0.25±0.00
14:0 1.83±0.21 1.48±0.59 1.81±0.23 3.79±0.31
15:0 0.83±0.51 0.67±0.30 0.40±0.08 0.46±0.03
16:0 21.90±0.01 21.21±0.91 24.53±0.01 22.83±2.12
17:0 2.15±0.06 2.24±0.14 1.76±0.31 1.35±0.09
18:0 10.86±0.56 12.98±2.99 13.17±0.87 9.28±0.95
Total saturated fatty acids 37.72±0.45 38.76±1.34 41.80±0.88 37.96±3.52

14:In5 0.13±0.07 0.15±0.01 0.18±0.01 0.32±0.00

iso 15:00 0.62±0.78 0.47±0.21 0.39±0.07 0.29±0.00
16:In7 7.14±0.25 5.59±3.57 4.12±1.23 7.59±0.20
iso 17:00 0.83±0.31 0.77±0.25 0.63±0.09 0.90±0.02
α 1so 17:00 0.79±0.02 0.77±0.11 0.45±0.02 0.45±0.01
17:1n9 1.43±0.02 1.43±0.28 0.87±0.02 0.96±0.07
18:1n9 9.67±0.15 9.24±0.70 12.05±1.48 15.73±1.21
18:1n7 6.12±0.03 5.30±1.44 5.15±0.06 5.24±0.43
20:1n9 0.57±0.14 0.69±0.06 0.00±0.00 0.50±0.03
Total monounsaturated fatty acids 27.30±0.04 24.41±6.47 23.84±2.92 31.98±1.94

18:2n6 1.7±0.02 1.24±0.51 2.97±0.25 3.83±0.20

18:2n4 1.86±0.46 1.30±1.54 0.00±0.00 0.43±0.31
18:3n6 0.45±0.78 0.44±0.20 0.00±0.00 5.49±7.49
18:3n3 2.10±0.27 1.36±1.07 2.63±0.46 4.64±0.24
20:3n6 1.05±1.04 1.26±0.12 0.32±0.01 0.00±0.96
20:4n6 5.91±0.49 7.32±1.71 6.87±0.68 3.24±0.27
20:5n3 (EPA) 4.95±1.05 3.89±1.99 0.77±0.03 1.66±0.18
21:5n3 0.95±0.14 1.37±0.51 0.56±0.02 0.25±0.02
22.4n6 0.85±0.59 1.44±0.86 0.97±0.04 0.52±0.03
22:5n6 1.16±1.07 2.24±1.29 1.05±0.04 0.40±0.06
22:5n3 3.97±0.37 3.65±0.20 3.80±0.83 3.17±0.09
22:6n3 (DHA) 7.41±0.47 7.36±0.87 12.26±1.46 5.64±0.75
DHA/EPA 1.50 1.89 15.92 3.40
Total polyunsaturated fatty acids 32.36±0.08 32.87±2.32 32.20±2.13 29.27±5.50

C1 0.49±0.24 1.12±0.76 0.66±0.02 0.28±0.02

C2 2.13±0.38 0.00 1.49±0.05 0.54±0.03
Total cholesterol 2.62±0.25 1.12 2.15 0.82

(n - 3) total 19.38 17.63 20.02 15.36

(n - 6) total 11.12 13.94 12.18 13.48
n-3/n-6 1.74 1.26 1.64 1.14

L. niloticus analysed. Further results from the investigations (Figure 1). There were, however, variations; for example,
indicated that all the portions of L. niloticus contained the abundance of 16:0 increased with fish length. On the
high quantities of 16:0, 18:0, 16:1n7, 18:1n9, 18:1n7, other hand, the abundance of 16:1n7, 18:1n9, 18:2n6
18:3n3, 18:2n6, 20:4n6, 20:5n3, 22:5n3 and 22:6n3 and 18:3n3 in the eggs was higher than that in fish of 0 to
 Namulawa et al. 131

Table 3. Fatty acid content (expressed as percentage of total fatty acids) in the caudal muscle of L. niloticus.

Fish size classes (cm)

Caudal
0-10 11-20 21-30 31-40
Fatty acid N=3 N=3 N=3 N=3
12:0 0.00±0.00 0.00±0.00 0.00±0.00 0.25±0.14
14:0 0.91±0.03 0.97±0.03 0.76±0.13 3.10±0.73
15:0 0.50±0.07 0.49±0.02 0.32±0.56 0.39±0.03
16:0 19.29±0.77 22.2±0.10 21.54±0.89 23.75±0.86
17:0 1.85±0.09 2.04±0.05 1.15±1.89 1.03±0.01
18:0 13.02±0.08 11.47±0.11 12.12±0.79 9.35±0.58
Total saturated fatty acids 35.57±0.72 37.17±0.27 35.89±0.35 37.87±1.19

14:In5 0.00±0.00 0.09±0.00 0.00±0.00 0.16±0.01

iso 15:00 0.27±0.00 0.51±0.02 0.32±0.47 0.27±0.11
16:In7 3.32±0.14 4.77±0.22 2.44±1.47 7.43±1.69
iso 17:00 0.75±0.08 0.67±0.02 0.57±0.29 0.82±0.14
α 1so 17:00 0.61±0.01 0.53±0.21 0.44±0.71 0.45±0.05
17:1n9 1.00±0.04 1.25±0.44 0.84±0.83 0.85±0.06
18:1n9 8.16±0.83 8.80±0.58 8.62±0.96 12.86±0.58
18:1n7 4.75±0.88 6.36±0.30 4.66±1.89 5.39±0.20
20:1n9 0.55±0.03 0.75±0.03 0.84±0.19 1.77±0.09
Total monounsaturated fatty acids 19.41±0.36 23.73±1.22 18.73±0.25 30.00±2.95

18:2n6 1.37±0.26 1.67±0.05 2.27±0.15 2.72±0.17

18:2n4 2.33±0.13 0.00±0.00 2.24±0.16 1.94±0.10
18:3n6 0.00±0.00 0.00±0.00 0.00±0.00 0.00±0.00
18:3n3 1.13±0.03 1.60±0.30 1.50±1.47 3.44±0.26
20:3n6 0.96±0.79 1.53±0.05 1.20±0.68 1.72±0.09
20:4n6 6.62±0.06 7.12±0.04 8.66±1.74 4.91±1.78
20:5n3 (EPA) 2.97±0.80 6.18±0.86 2.92±1.79 1.88±0.53
21:5n3 0.00±0.00 0.97±0.03 0.62±1.69 0.00±0.00
22.4n6 0.88±0.05 0.54±0.02 1.02±0.59 1.05±0.05
22:5n6 1.83±0.69 1.41±0.05 1.92±0.48 0.75±0.04
22:5n3 3.49±0.68 5.48±0.21 4.31±0.39 4.29±0.21
22:6n3 (DHA) 19.41±1.75 10.29±0.43 15.23±1.69 7.47±2.40
DHA/EPA 6.54 1.67 5.22 3.97
Total polyunsaturated fatty acids 40.99±1.78 36.79±1.04 41.89±0.25 30.17±4.21

C1 1.15±0.60 0.69±0.02 1.02±0.49 0.59±0.03

C2 2.88±0.82 1.59±0.06 2.58±1.59 1.37±0.07
Total cholesterol 4.03±1.22 2.28±0.64 3.60±1.10 1.96±0.55

(n - 3) total 27.00 20.46 24.58 17.08

(n - 6) total 11.66 9.41 15.07 11.15
n-3/n-6 2.32 2.17 1.63 1.53

40 cm total length. Table 1 also shows that cholesterol correlation (r2 = 0.715, p<0.0001, 2-tailed, α = 0.05),
isomers were detected in the abdominal and caudal between the composition of fatty acids in the eggs and in
muscles but not in the eggs or the egg fat. the egg fat. There was however little variance (p<0.05)
Statistical analysis indicated that there was a significant between the mean fatty acid percentage in the various
132 Afr. J. Pure Appl. Chem.

35
Eggs

30 0 to cm
10 cm
0-10

25 11 to 20 cm
Fatty acid (%)

20 21 to 30 cm

31 to 40 cm
15

10

0
7

3
0

n7

n
:0

:0

:In
:1

:1

:3

:2

:4

:5

:5

:6
16

18

18

18

18

18

20

20

22

22
16

Fatty acids

Figure 1. Abundance of fatty acids in the L. niloticus eggs and fish of 0 to 40 cm (TL).

class sizes of the abdominal and caudal muscles. 0.30 cm length before changing to insect larvae and
eventually to a fish diet (haplochromines, tilapianes,
Barbus, among others) on attaining a total length of 2.0
DISCUSSION cm (Hopson, 1972). This could explain why L. niloticus
juveniles tend to be restricted around shallow vegetated
Fatty acid composition in animals depends on a number areas were most zooplankton is found (Ogutu-Ohwayo,
of factors such as sex, age, size and body temperature. 1985). Much as the mouth gape determines the size of
As temperature decreases, the level of unsaturation the prey that can be swallowed whole by a predator (De
tends to increase for fish so as to maintain membrane Silva and Anderson, 1995; Lovell, 1998; Halver and
fluidity and general body flexibility. When the surrounding Hardy, 2002), L. niloticus rapidly moves up the trophic
temperatures increase, an increase of phospholipids is level presumably due to the need to satisfy the various
necessary to counter excessive fluidity (Eastman, 1990; fatty acid requirements at different growth stages.
Martino and Cruz, 2004). This phenomenon seems to In consonance, there was significant difference
work well in fish species that experience extreme weather (p<0.005) between the mean fatty acid percentage in the
conditions especially in bitterly cold winters and very hot eggs and at the 0 to 10 cm length stage. Figure 1 shows
summers. This was not observed in L. niloticus where the that the fatty acids: 16:0, 18:0, 16:1n7, 18:1n9, 18:1n7,
concentration of polyunsaturated fatty acids (22.16±0.81 18:3n3, 18:2n6, 20:4n6, 20:5n3, 22:5n3 and 22:6n3 are
to 43.64±14.32%), monounsaturated fatty acids particularly abundant in L. niloticus of 0 to 40 cm length,
(18.73±0.25 to 42.98±11.8%) and saturated fatty acids which is indicative of these acids possibly being required
(13.39±2.45% to 40.18±0.48%) is almost similar, owing to in the constitution of the eggs, larvae and juveniles
the species being a tropical fish that does not experience (Izquierdo et al., 2001). The abundances of the fatty
such extreme climatic conditions. Age impacts acids 16:1n7, 18:1n9, 18:2n6 and 18:3n3 in the eggs is
significantly on the variation of fatty acid dietary high compared to that in the 0 to 40 cm fish, implying a
requirements in fish (Izquierdo and Fernandez-Palacios, particular necessity for these acids in the eggs of L.
1997; Kolkovski, 2005). Fish exhibit a variety of food niloticus. Docosahexaenoic (22:6n3), palmitic (16:0),
intake at their different stages of development. In this eicosapentaenoic (20:5n3) and oleic (18:1n9) acids have
respect, L. niloticus feeds on zooplankton (Cladocera) at been found to be of particular importance in the egg
 Namulawa et al. 133

constitution of halibut (Hippoglossus hippoglossus), improving maturation and spawning quality (Sorbera et
turbot (Scophthalmus maximus), plaice (Pleuronectes al., 2001). Docosahexaenoic is also necessary for
platessa), dolphin (Coryphena hyppurus), red sea bream reproduction, survival and disease prevention (Izquierdo,
(Pagrus major) and gilthead seabream (Sparus aurata) 2000). The high docosahexaenoic/eicosapentaenoic
(Izquierdo et al., 2001). Investigations performed by ratios in the L. niloticus eggs and egg fat (7.38 and 3.86,
Ugoala et al. (2009) indicate that fatty acids: 16:0, 18:0, respectively) could be used to account for the need of
18:1n9, 18:1n7, 18:3n3, 20:4n6 and 20:5n3 were similarly this fatty acid for egg and post-hatching development.
important in L. niloticus from Kainji Lake dam site in Due to their importance in fish development, the three
Nigeria; however, the fatty acids 16:1n7, 18:2n-6, 22:5n3 polyunsaturated fatty acids: docosahexaenoic,
and 22:6n3 were not detected. This could imply that fatty eicosapentaenoic and arachidonic are considered
acid requirements are greatly affected by the surrounding essential for freshwater species (Yu and Sinhuber, 1975).
environment, given that Kainji Lake dam site is located Inadequate supply of these essential fatty acids in the
farther away from the equator compared to Lake Victoria. diet gives rise to poor feeding and swimming activities,
Similar changes in fatty acid composition have been poor growth, fatty liver, hydops (increased water content
observed in Oreochromis niloticus and Clarias gariepinus of the muscle), shock syndrome, fin erosion,
(Ugoala et al., 2008), probably because of the more mitochondrial swelling, haemoglobin deficiency, swim
vigorous carnivorous nature of the latter compared to that bladder inflation, abnormal pigmentation, disaggregation
of the former. of gill epithelia, immune deficiency and raised cortisol
Docosahexaenoic acid has also been found to levels (Izquierdo, 1997). Inappropriate quantities of
accumulate in the larval and juvenile fish stages of essential fatty acids in broodstock diets also lead to
Sparus aurata, Clupea harengus and Scophthalmus reduced fecundity and fertilisation rates, embryo
maximus (Mourente, 2003). It has been suggested that deformity and damaged larval quality (Izquierdo et al.,
these fatty acids are of vital importance to the further 2001). It would therefore be essential to include these
development of the embryos and larvae of the common critical quantities of the essential fatty acids in artificial L.
snook (Yanes-Roca et al., 2009). This could explain the niloticus larval and broodstock diets. One major
critical concentrations of these fatty acids in the Nile difference is observed between the essential fatty acid
perch eggs (Docosahexaenoic: 8.95±1.57%, palmitic: requirements of fresh and those of marine-water fish
8.37%±8.37, eicosapentaenoic: 1.04±0.16% and oleic: (Ugoala et al., 2008); the former require either linoleic
3.53±1.13%). This observation points to the fact that any acid (18:2n-6) or linolenic acid (18:3n3) or both, while for
artificial feed formulated for purposes of viable Nile perch the latter the highly unsaturated fatty acids:
broodstock and larval development should contain these eicosapentaenoic (20:5n3) and/or docosahexaenoic
fatty acids in this range of composition. Similar studies (22:6n3) are necessary (Sargent et al., 1999). Several
carried out on fatty acid profiles of fish (Izquierdo et al., authors (Ackman et al., 2002; Tocher, 2003; González et
2003; Tocher and Dick, 2004; Osman et al., 2007; al., 2006) have suggested that fresh-water fish contain
Ugoala, 2008; Ugoala et al., 2009) indicate that fatty high concentrations of 20:4n6 and 18:2n6 fatty acids, as
acids are vital in several metabolic activities. compared to their marine counterparts. This may be
Eicosapentaenoic, docosahexaenoic and arachidonic attributed to the diet of some fresh-water fish which
acids are essential for growth, development and survival includes algae, insects and insect larvae. Unlike other
(Sargent et al., 1999). Eicosapentaenoic and herbivorous fresh-water fishes, the diet of L. niloticus
docosahexaenoic acids in particular are heavily involved comprises mainly tilapines and haplochromines that in
in the development of the nervous system, including the turn depend on freshwater algae (Fraser et al., 1989).
brain and visual cells (Furuita et al., 1998; Cahu et al., These algae are rich in linoleic and linolenic acids
2003). This could be responsible for the relatively high which are responsible for the remarkably high levels of
levels of docosahexaenoic (8.95±1.57% in the eggs and 20:4n6 fatty acids; this ties up very well with the
7.41±0.47% in 0 to 10 cm juveniles) in L. niloticus substantial amounts of the fatty acid 20:4n6 found in L.
because the species needs this fatty acid for visual niloticus in this work. Fresh-water fish fatty acids exhibit
development, owing to L. niloticus being a vigilant visual the rather unusual ability to elongate their carbon chains
feeder (Hamba, 2002). Eicosapentaenoic and through enzymatic desaturation (Steffens, 1997). This
docosahexaenoic acids also improve vitality and stress observation is contrary to what is observed in the fatty
resistance (Watanabe and Kiron, 1994), so that the acids found in other strictly carnivorous fish species
dietary inclusion of these acids in the levels observed in (Gutierrez and Da Silva, 1993; Jobling, 2004; Halilo,
the eggs (1.04±0.16% and 8.95±1.57%, respectively) and 2004). Fatty acids are indispensable in the fish diet
larvae (4.97±1.05% and 7.41±0.47%, respectively) will because of their numerous functions in the metabolism
probably be of great significance for the survival and and functioning of the body; yet, owing to their restricted
visual development of Nile perch larvae. mobility, farmed fish can only access such fatty acids
On the other hand, arachidonic acid is important for through the feeds provided. It is therefore important that
134 Afr. J. Pure Appl. Chem.

fish feed formulators make appropriate fatty acid González S, Flick GJ, O’Keefe SF, Duncan SE, McLean E, Craig SR
(2006). Composition of farmed and wild yellow perch (Perca
inclusions in the fish diets so that the fish can attain flavescens). J. Food Compos. Anal., 19: 720–726.
optimal growth potential without fatty-acid related Grahl-Nielsen O, Barnung T (1985). Variations in the fatty acid profile of
deficiencies. marine animals caused by environmental and developmental
changes. Mar. Environ. Res., 17: 218-221.
Gregory RG (2006). The Nile perch lates niloticus: a potential candidate
for cage aquaculture. In M. Halwart JF (eds). FAO Regional
Conclusion Technical Expert Workshop on cage culture in Africa. Entebbe
Uganda, 20-23 October 2004. FAO Fish. Proc. Rome, FAO. 6: 11.
The presence of twenty-seven different fatty acids have Gutierrez LE, da Silva RCM (1993). Fatty acid composition of
been confirmed in L. niloticus, most of which have been commercial fish from Brazil. Sci. Agric. Piracicaba, 50: 478-483.
Halilo HL, Bayır A, Sirkecio AN, Mevlüt NA, Atamanalp M (2004).
attributed to the species being a tropical fish that does Comparison of fatty acid composition in some tissues of rainbow trout
not experience extreme climatic conditions. (Oncorhynchus mykiss) living in seawater and freshwater. Food
Docosahexaenoic acid content in L. niloticus is relatively Chem., 86: 55-59.
Halver J, Hardy RW (2002). Fish nutrition. Academic Press, London, p.
high possibly due to this fatty acid being of vital 824.
importance in young fish. It would consequently be Hamba S (2002). Observations from initial attempts to culture Nile
necessary in any subsequent formulation of feeds for perch. Diploma project report. Fisheries Training Institute. Entebbe,
cultured L. niloticus to take into consideration the Uganda.
Hopson AJ (1972). The study of the Nile perch (Lates niloticus) (Pisces:
inclusion of 16:0, 18:0, 16:1n7, 18:1n9, 18:1n7, 18:3n3,
Centropomidae) in Lake Chad. Overseas Research Publication,
18:2n6, 20:4n6, 20:5n3, 22:5n3 and 22:6n3 fatty acids in Commonwealth Office, Overseas Development Administration, Her
the broodstock, larval and juvenile diet. Majesty’s Stationery Office, 19: 93.
Izquierdo M (2000). Essential fatty acid requirements in Mediterranean
fish species. Cahiers Options Méditerranéennes, 63: 91-102.
Izquierdo M, Fernandez-Palacios H, Tacon AGJ (2001). Effect of
ACKNOWLEDGEMENTS broodstock nutrition on reproductive performance of fish.
Aquaculture, 197: 25-42.
The authors would like to express their gratitude to the Izquierdo M, Fernandez-Palacios H (1997). Nutritional requirements of
World Bank, through the millennium science initiatives marine fish larvae and broodstock. In Tacon A, Basurco B, eds.
Feeding Tomorrow’s Fish, pp. 243-264.
(MSI) and the Uganda National Council for Science Izquierdo M, Obach A, Arantzamendi L, Montero D, Robaina L,
Technology (UNCST), for financial support. Our sincere Rosenlund G (2003). Dietary lipid sources for seabream and
thanks also go to the anonymous reviewer for his/her seabass: Growth performance, tissue composition and flesh quality.
very constructive comments that led to a marked Aquacult. Nutr., 9: 397.
Jobling M (2004). ‘Finishing’feeds for carnivorous fish and fatty acid
improvement in this manuscript. dilution model. Aquacult. Res., 35: 706-709.
Kolkovski S (2005). Progress in fish larval nutrition live feed enrichment
and brood stock additives - Tailor made to requirements. Aqua
REFERENCES Feeds: Formulation Beyond, 2: 7-8.
Lake Victoria Fisheries Organization (LVFO) (2009). Nile perch Market
Abbas KA, Mohamed A, Jamilah B (2009). Fatty acids in fish and beef Report, 2009.
and their nutritional values: A review. WFL Publisher Science and Lovell RT (1998). Nutrition and feeding of fishes. Chapman and Hall,
Technology. J. Food Agric. Environ., 7: 3-4. New York.
Ackman RG, McLeod C, Rakshit S, Misra KK (2002). Lipids and fatty Martino RC, Cruz GM (2004). Proximate composition and fatty acid
acids of five freshwater food fishes of India. J. Food Lipids, 9: 127- content of the mangrove oyster Crassostrea rhizophorae along the
145. year seasons. Braz. Arch. Biol. Techn., 47: 955-960.
Bwanika GN, Chapman LJ, Kizito Y, Balirwa J (2006). Cascading Mourente G (2003). Accumulation of DHA (docosahexaenoic acid;
effects of introduced Nile perch (Lates niloticus) on the foraging 22:6n-3) in larval and juvenile fish brain. In the Big Fish Bang.
th
ecology of Nile tilapia (Oreochromis niloticus). Ecol. Freshwater Fish, Proceedings on the 26 Annual Larval Fish conference, (Eds)
15: 470-481. Howard J, Brownman and Anne Berit. Institute of Marine Research,
Cahu C, Zambonino JI, Takeuchi T (2003). Nutritional components Norway.
affecting skeletal development in fish larvae. Aquaculture, 227: 254- Mourente G, Tocher DR (1992a). Lipid class and fatty acid composition
258. of brain lipids from Atlantic herring (Clupea harengus, L) at different
De Silva SS, Anderson TA (1995). Fish nutrition in aquaculture. stages of development. Mar. Biol., 112: 553-558.
Chapman and Hall Aquaculture series, London. Ogutu–Ohwayo R (1985). The effects of predation by Nile perch
Eastman JT (1990). The biology and physiological ecology of introduced in Lake Kyoga in relation to the fisheries of Lake Kyoga
notothenioid fishes. In Gon O and Heemstra PC (eds.,) Fishes of and Victoria. FAO Fish. Rep., 355: 18-41.
Sothern Ocean. JLB Smith Institute of Ichthyology. Grahamstown, pp. Ogwok P, Muyonga JH, Sserunjogi ML (2008). Fatty acid profile and
34-51. stability of oil from belly flaps of Nile perch (Lates niloticus). Food
Fraser AJ, Sargent JR, Gamble JC, Seaton DD (1989). Formation and Chem., 108: 103-109.
transformation of fatty acids in an enclosed marine food chain Okedi JO (2005). A Historical perspective of the challenges and
comprising phytoplankton, zooplankton and herring (Clupea opportunities in management of the Lake Victoria fisheries: Lessons
harengus L) larvae. Mar. Chem., 27: 1-18. for the future. Proceedings of the Regional Stakeholders’
th th
Furuita H, Takeuchi T, Uematsu K (1998). Effects of eicosapentaenoic Conference, Imperial Resort Beach Hotel, Entebbe 24 -25
and docosahexaenoic acids on growth, survival and brain February, 2005.
development of larval Japanes flounder (Paralichtys olivaceus). Osman IJ, Khaza’ai H, Ridzwan H (2007). Fatty acid profiles of fin fish
Aquaculture, 161: 269-279. in Langkawi island, Malaysia. J. Oleo. Sci., 56: 107-113.
 Namulawa et al. 135

Rimmer MA, Reed A (1989). Effects of nutritional enhancement of live Turon F, Rwabwogo B, Baréa B, Pina M, Graille J (2005). Fatty acid
food organisms on growth and survival of Barramundi/Seabass Lates composition of oil extracted from Nile perch (Lates niloticus) head. J.
calcarifer (Bloch) larvae. Adv. Trop. Aquacult., 9: 611-623. Food Compos. Anal., 18: 717-722.
Sargent JR, Bell JG, Henderson RJ, Tocher DR (1995). Requirement Ugoala C, Ndukwe GI , Audu TO (2008). Comparison of Fatty Acids
criteria for essential fatty acids. J. Appl. Ichthyol., 11: 183-198. profile of some freshwater and marine fishes. Internet J. Food Saf.,
Sargent JR, McEvoy L, Estevez A, Bell G, Henderson J, Tocher D 10: 9-17.
(1999). Lipid nutrition of marine fish during early development: Ugoala C, Ndukwe GI, Audu TO (2009). Fatty acids composition and
Current status and future directions. Aquaculture, 179: 217-229. nutritional quality of some freshwater fishes. Nature Precedings: doi:
Sorbera LA, Asturiano JF, Carrillo M, Zanuy S (2001). Effects of 10.1038/npr.2009.3239.1: posted 12 May, 2009.
polyunsaturated fatty acids and prostaglandins on oocyte maturation Uotani I, Saito T, Hiranuma K, Nishikawa Y (1990). Feeding habit of
in a marine teleost, the European sea bass (Dicentrarchus labrax). bluefin tune Thunnus thynnus larvae in the western North Pacific
Biol. Reprod., 64: 382-389. Ocean. Nippon Suisan Gakkaishi, 56: 713-717.
Steffens W (1997). Effects of variation in essential fatty acids in fish Watanabe T, Kiron V (1994). Prospects in larval fish dietetics.
feeds on nutritive value of freshwater fish for humans. Aquaculture, Aquaculture, 124: 223-253.
119: 63-88. Yanes-Roca C, Rhody N, Nystrom M , Main KL (2009). Effects of fatty
Tang H, Chen L, Xiao C, Wu T (2009). Fatty acid profiles of muscle acid composition and spawning season patterns on egg quality and
from large yellow croaker (Pseudosciaena crocea R.) of different age. larval survival in common snook (Centropomus undecimalis)
J. Zhejiang Univ. Sci., 10: 154-158. Aquaculture, 287: 335-340.
Tocher DR (2003). Metabolism and function of lipids and fatty acids in Yu TC, Sinnhuber RO (1975). Effect of dietary linolenic and linoleic
teleost fish. Rev. Fish Sci., 11: 107-184. acids upon growth and lipoid metabolism of rainbow trout (Salmo
Tocher DR, Dick JR (2004). Essential fatty acid deficiency in freshwater gairdneri). Lipids, 10: 63-66.
fish: The effects of linoleic, α-linolenic, γ-linolenic and stearidonic
acids on metabolism of [1-14C] 18:3n-3 in a carp cell model. Fish
Physiol. Biochem., 22: 67-75.