2006 The Authors

Journal compilation  2006 Blackwell Munksgaard

Ecology of Freshwater Fish 2006: 15: 441445

Printed in Singapore All rights reserved

ECOLOGY OF
FRESHWATER FISH

Variations in fatty acids composition of pikeperch

(Sander lucioperca) liver with respect to gonad
maturation
Uysal K, Yerlikaya A, Aksoylar MY, Yontem M, Ulupinar M. Variations in
fatty acids composition of pikeperch (Sander lucioperca) liver with respect
to gonad maturation.
Ecology of Freshwater Fish 2006: 15: 441445.  2006 The Authors.
Journal compilation  2006 Blackwell Munksgaard
Abstract We examined seasonal variations in liver condition factor, total
lipid and fatty acid composition of maturing pikeperch (Sander lucioperca)
from Lake Egirdir, Turkey. The spawning of pikeperch in Egirdir Lake
started in March, continued very fast during May and gradually ended in
July as determined by measuring the gonadosomatic index. The total lipid
content of male and female liver reached its highest level in September and
November (the end of dense feeding period during which temperature
decreased sharply), but declined to the lowest level in May (just after
spawning). The highest level of liver condition factor was observed in
January and March during which gonads mature rapidly. Palmitic acid was
the predominant saturated fatty acid (SFA) in female and male liver. The
ratio of monounsaturated fatty acids (MUFA) was higher than that of
polyunsaturated fatty acids (PUFA) in both sexes. Docosahexaenoic acid
(22:6n-3), eicosopentaenoic acid (20:5n-3) and arachidonic acid (20:4n-6)
were the predominant PUFA. Seasonal variations in PUFA were more
evident than that in other fatty acids. PUFA, especially the n-3 fatty acids
ratio, increased to the maximum level during the sharp decreases in
temperature (in November). However, the amount of PUFA decreased to
the lowest level in reproductive period. The results suggest that the ratio of
pikeperch liver fatty acids is inuenced by gonad maturation and
temperature variations and that pikeperch requires a large amount of PUFA
and the n-3 fatty acids for the development of gonads.

Introduction

Pikeperch (Sander lucioperca), a species of European

origin, was rst introduced into Lake Egirdir in 1955.
Since then, it has changed the biological balance of
the lake and has become the dominant sh species in
the lake. Prior to the introduction of this species, it
was recorded that there were 10 different sh species
in the lake (Sarhan 1970). Early commercial sheries
were based on the species of Cyprinus carpio,
doi: 10.1111/j.1600-0633.2006.00174.x

K. Uysal1, A. Yerlikaya1,
M. Y. Aksoylar2, M. Yontem1,
M. Ulupinar3
1

Department of Biology, Faculty of Art &

Sciences, Dumlupinar University, Kutahya,
Turkey, 2Faculty of Water Products, University of
Suleyman Demirel, Isparta, Turkey, 3Water
Resources Department, Ministry of Agriculture,
Ankara, Turkey

Key words: Egirdir Lake; unsaturated fatty acids;

spawning; gonad maturation; seasonal variations
K. Uysal, Department of Biology, Faculty of Art &
Sciences, Dumlupinar University, Kutahya,
Turkey; e-mail: kuysal@dumlupinar.edu.tr
Accepted for publication April 12, 2006

Acanthorutilus handlirschi and Varicorhinus pestai.

Eight of these species had disappeared from the lake
by the beginning of 1990s. While annual total sh
catch was 622 ton in 1979, this value decreased to
145 ton in 1990. To improve sheries in the lake, rst
Carassius auratus gibelio at the beginning of the
1990s and later Tinca tinca in 1997 were introduced
into the lake. Of these two species, only C. auratus
gibelio population increased rapidly, and in the
following several years it has become the dominant
441

Uysal et al.
sh in the lake (Balk 1999; Balk et al. 2004; Celik
et al. 2005).
Pikeperch, which is distributed abundantly in
freshwater in Turkey and has commercial value in
both domestic and foreign markets, is a lean freshwater sh. To our best knowledge, no reports have yet
been published about the effects of sexual and
seasonal variations on the liver fatty acid composition
of pikeperch.
The total lipid and lipid fatty acid composition of all
sh species varies according to the season, geographical location of the catch, size, sex and the state of their
reproductive cycle (Cai & Curtis 1990; Shirai et al.
2001, 2002; Luzia et al. 2003). While the fatty shes
(e.g. salmon, trout and herring) store fats predominantly in muscle tissue, the lean shes deposit their fats
in body cavities and perivisceral organs (e.g. liver).
Therefore, the livers of lean shes are commonly more
fatty than fatty shes. Although the muscle of Gadus
morhua (cod) contain insignicantly small amount of
fat, the liver is composed of approximately 6070% of
lipid (Jangaard et al. 1967; Addison et al. 1968). In
many spring-spawning shes, gonads mature during
winter when food supplies are limited and feeding is
reduced. Therefore, nutrients for germinal tissue
growth have to be drawn from other tissues like
muscle and liver. It was reported that females use
mainly saturated fatty acids (SFA) for meeting the
energy requirement of gonad maturation while males
use monounsaturated fatty acids (MUFA). However,
during vitellogenesis, females can also mobilise stored
precursors (e.g. linoleic and linolenic acids) of fatty
acids of n-3 and n-6 family for gonad maturation
(Medford & Mackay 1978; Cejas et al. 2003).
Fish liver oil (particularly cod liver oil) has long
been a preferred source of oil for the prevention of
vision and growth problems. There is a growing
demand for sh oils for the use in human food and
pharmaceuticals. The identication of suitable
sources rich in n-3 fatty acids to meet these demands
is thus very important (Jayasinghe et al. 2003). The
present study was performed to determine
the seasonal variations (especially with respect to
the gonad maturation, spawning and environmental
temperature) in liver fatty acid composition of
pikeperch.
Materials and methods
Study area

Lake Egirdir (37503857N, 30443057E) is

situated in the lakes district of Turkey. It is the second
largest natural freshwater lake in Turkey with a surface
area varing between 440 and 490 km2 and an altitude
of 914 m above sea level. The maximum depth of the
442

lake is 810 m. It is a tectonic lake with sand and

gravels in shallow places and bedrocks in some places.
The lake is mostly fed with spring waters from the lake
bottom. In the east shoreline, canes and reed beds are
abundantly found, whereas these are rarely seen on
other shorelines. The surface is partly covered with ice
in winter. In summer, the water temperature reaches up
to 30 C. The lake has an oligotrophic character
(Campell 1992; Becer 1995).
Pikeperches used in this study were caught using a
gill net and transported on ice to the laboratory. The
ages were determined from scales after biometric data
analyses (i.e. weight and length). All shes used in the
analyses were almost similar in size and ages (over
2 years old), and had also spawned the previous year
as determined by gross examination of gonad morphology.
Liver condition factor (KL) was determined by the
following standard equation: KL (total liver
weight/total sh weight) 100 (Jangaard et al.
1967). Gonadosomatic Index (GSI) values were
calculated according to the equation, GSI (total
weight of gonad/total body weight) 100 (Treasurer
1990). Sampling was performed once every
2 months. The liver from each sex was extracted
for analyses by homogenisation in a warring blender
for 3 min. A mixture of chloroform and methanol
(2:1) was used as a solvent. Total lipid extraction
was performed according to the procedure of Folch
et al. (1956). Lipids were trans-esteried with
Bf3methanol (Morrison & Smith 1964; Moss et al.
1974). Routine analysis of the methyl esters was
carried out on a Shimadzu GC 14B model equipped
with ame ionisation detector (FID). The chromatographic conditions were as follows: injection-port
temperature was at 225 C; detector temperature at
225 C; initial oven temperature at 135 C; but rising
to 185 C at a rate of 3 C min)1. The carrier gas was
nitrogen, set at a ow rate of 30 ml min)1. Samples
were injected into a 2.1 m 53-mm columns packed
with 10% DMCS on Chromosorb PAW-DMCS
(Sigma, St. Louis, MO, USA). The gas chromatography (GC) electrometer was connected to a Shimadzu
CR electronic integrator for automatic quantitative
calculations and programmed peak processing. The
peaks were identied by comparing the relative
retention time with pure methyl ester peaks (Sigma).
The results are reported as means SEM of 315
determinations. The statistical differences were analysed with one-way anova and the Duncan post-test.
Results were considered signicant at P < 0.05.
Results

The maturation of ovaries began in July and continued

rapidly in winter, and was completed in March. From

Seasonal variations in pikeperch liver fatty acids

6
Male
Female

% GSI

Nov

Sep

Jul

May

Mar

Jan

Fig. 1. Gonadosomatic index (GSI). GSI values were plotted as

means SEM (N 15).

2.4

Female condition factor

1.2

Male total lipid

5
0.6

Total lipid
(% of wet weight)

Condition factor

Male condition factor

1.8

Female total lipid

Nov

Sep

Jul

May

Mar

Jan

0.0

Fig. 2. Liver condition factor and total liver lipid. Liver condition
factor was determined as described by Jangaard et al. (1967).
Results are plotted as means SEM (N 15). The total lipid level
was determined by the procedure of Folch et al. (1956) using gas
chromatography. The results are graphed as means SEM (N 3
at each data point).

July to March, a ninefold increase in GSI was

observed. On the other hand, the maturation of testis
started to increase in September and levelled off in
November (with a sixfold increase in GSI between
these 2 months). The spawning period of pikeperch
began in March, continued signicantly during May
and gradually completed in July as determined by the
GSI value (Fig. 1). The highest liver condition factors
of both sexes were observed in January, but reached its
minimum level in September and November (Fig. 2).
Total lipid content of male and female liver was about
5% of wet weight in March, but it increased to 7% of
wet weight (the highest amount observed) in November (Fig. 2). While statistically signicant differences
(P < 0.05) in GSI values between male and female
shes were recorded, no signicant changes in
condition factor or total lipid content of livers were
detected between sexes. Major fatty acids and their

seasonal variations in pikeperch liver lipids are

indicated in Table 1. SFA changed annually between
28% and 45% in male liver and between 31% and 42%
in female liver. Palmitic acid (16:0) was the predominant SFA, accounting for approximately 7080% of
the total SFA. Unsaturated fatty acids (UFA) constituted more than half of the total fatty acids in both
female and male liver. The ratio of MUFA was found
to be higher than that of polyunsaturated fatty acids
(PUFA). Oleic acid (18:1) was the most abundant
MUFA. Eicosapentaenoic acid (20:5n-3), docosahexsaenoic acid (22:6n-3) and arachidonic acid
(20:4n-6) were the predominant PUFA. The n-3 PUFA
were present in excess of n-6 PUFA in both male and
female liver. The ratios of SFA and UFA in liver were
not signicantly different between female and male
shes (P > 0.05). Nonetheless, signicant differences
between female and male MUFA (especially oleic
acid) were observed in the current study. Likewise,
signicant differences were detected when female
PUFA were compared with male PUFA (Table 1).
The relative proportion of SFA in liver changed
seasonally from 45% to 28% and from 42% to 31%
in male and female, respectively. The ratio of SFA in
fatty acid composition decreased to a minimum in
November. In this period, the water temperature
(9.8 C) signicantly decreases and gonad maturation
begins. On the other hand, the concentration of PUFA
in liver varied seasonally from 32% to 13% and from
28% to 15% in male and female, respectively
(Table 1). The relative concentration of n-3 polyenes
varied seasonally ranging from 19.5% to 5% and from
16% to 6% annually in male and female liver,
respectively. The relative proportion of PUFA in liver,
particularly n-3 fatty acids, diminished signicantly
with the maturation of gonads and increased during
dense feeding periods in summer, and reached its
maximum level in fall. The monthly proles of MUFA
were more similar to that of SFA rather than to that of
PUFA.
Discussion

The amount of lipids in liver was maximum in

September and November, which is a dense feeding
period. During dense feeding periods (i.e. September
and November), both liver condition factor and GSI
were signicantly lower as compared with the level in
other months, but were highest in January and March.
In light of this evidence, it is thought that the liver
condition of pikeperch may be associated with gonad
maturation rather than feeding, as it is known that
there is a decrease in feeding in January and March.
The reason for higher liver condition factor observed
in these months may be because of the temporal and
specic expression of factors in liver, which play
443

Uysal et al.
Table 1. Seasonal variations in male and female liver fatty acids of pikeperch (per cent of total fatty acids).
January
Male
C12:0
C14:0
C15:0
C16:0
C16:1
C18:0
C18:1
C18:2 n-6
C18:3 n-3
C20:1
C20:2 n-6
C20:4 n-6
C20:5 n-3
C22:6 n-3
C24:1
P
Other
P
SFA
P
UFA
P
MUFA
P
PUFA
P
n-3
P
n-6

0.03
1.6
0.6
35.6
3.1
2.6
22.3
2.0
1.4
2.4
1.3
4.2
6.2
7.0
2.0
7.1
40.5
52.2
30.0
22.2
14.6
7.5

March
Female

0.0
0.6
0.1
2.4
0.0
1.3
4.0 29.7
0.4
5.2
0.6
3.2
2.3 25.7
1.3
1.7
1.0
1.1
0.1
1.5
0.5
1.2
0.5
5.6
0.4
4.2
1.8
5.5
1.0
1.9
3.0
7.5
3.4 37.4
6.5 54. 0
3.7 34.4
3.9 19.6
2.0 10.9
2.4
8.7

May

Male
0.3
0.4
0.7
2.8
0.5
0.3
1.0
0.8
0.4
0.0
0.0
0.1
1.2
1.4
0.8
1.0
3.8
2.7
0.5
4.1
2.3
0.8

0.9
2.4
1.4
35.2
4.9
3.3
26.7
0.6
1.1
3.2
0.7
5.1
3.5
3.1
0.7
6.1
43.3
50.4
35.9
14.4
7.9
6.5

Female

0.8
0.5
0.3
1.8
0.3
0.5
3.2
0.4
0.8
0.4
0.3
1.6
0.6
0.8
0.0
1.7
1.0
1.0
2.5
2.9
1.8
1.2

0.2
1.4
0.5
33.5
2.2
3.0
30.2
2.1
2.0
1.3
1.2
6.5
3.1
3.5
0.6
7.1
38.9
53.2
34.5
18.6
8.6
9.9

July

Male
0.1
0.0
0.0
0.6
0.4
0.1
1.9
0.1
0.5
0.0
0.1
0.4
0.1
0.4
0.1
1.6
0.6
1.0
2.0
1.0
0.8
0.2

0.3
3.0
1.6
36.0
4.2
3.8
30.6
0.8
1.7
1.0
0.8
5.9
3.1
2.5
0.7
3.4
44.9
51.5
36.6
14.8
7.4
7.6

Female

0.1
0.7
0.4
2.5
0.4
0.5
3.1
0.4
0.8
0.2
0.2
1.0
0.6
1.1
0.0
0.2
1.4
1.3
3.2
4.0
2.4
1.6

0.1
2.5
0.6
33.1
2.6
4.3
27.4
2.1
2.4
2.0
1.7
5.2
2.2
1.6
0.7
9.9
40.8
48.2
32.8
15.3
6.2
9.1

September

Male
0.0
0.3
0.0
0.9
0.4
0.1
2.1
0.2
0.3
0.4
0.4
0.4
1.3
0.7
0.2
0.8
1.0
0.2
2.0
2.0
2.3
0.2

0.1
2.9
1.0
33.0
4.3
3.7
28.5
0.7
1.2
1.7
1.4
3.2
3.0
3.4
3.4
7.8
40.9
50.9
38.0
12.9
7.7
5.4

Female

0.0
0.1
0.1
1.8
0.2
0.3
1.9
0.4
0.8
0.5
0.7
0.2
0.4
0.1
1.5
1.1
2.0
1.0
1.2
1.8
0.6
1.1

0.7
1.5
1.5
32.1
1.3
3.6
26.8
2.5
2.2
1.4
1.4
5.3
6.9
4.3
1.5
6.0
39.3
54.5
31.1
23.4
14.5
9.2

Male
0.1
0.4
0.1
4.2
0.5
0.3
2.8
0.0
0.3
0.2
0.0
0.5
0.9
2.0
0.5
2.7
4.9
2.2
1.5
3.8
3.0
0.5

0.3
3.3
0.8
36.6
4.2
3.7
26.1
2.7
1.9
4.8
2.6
3.0
1.3
1.5
2.2
4.2
44.9
50.8
37.4
13.3
4.8
8.4

November
Female

0.0
0.2
0.0
3.0
0.6
0.2
1.7
0.2
0.4
0.4
0.5
0.6
0.1
0.6
0.8
2.9
3.1
1.4
1.6
1.8
1.1
0.8

0.4
3.5
1.5
32.4
2.0
3.7
24.0
2.0
3.6
1.0
2.2
4.5
6.7
3.9
1.0
7.8
41.6
50.2
27.0
23.1
14.3
8.7

Male
0.1
0.0
0.0
0.8
0.6
0.1
1.1
0.4
0.0
0.2
0.4
0.0
0.0
0.1
0.3
2.6
0.8
1.8
1.8
0.1
0.1
0.7

0.05
1.1
1.8
21.1
2.3
4.1
17.1
2.0
2.3
2.3
1.1
9.4
9.2
7.7
2.7
14.7
28.3
57.0
24.6
32.3
19.5
12.6

Female

0.0
0.0
0.1
0.6
0.5
0.1
0.7
0.3
0.2
0.3
0.2
0.5
0.4
1.9
0.4
1.2
0.8
1.8
0.5
1.5
2.0
0.6

0.2
0.9
0.7
25.0
2.1
4.2
24.4
2.6
3.6
1.2
1.3
8.3
7.9
4.0
1.8
10.1
31.2
57.7
29.8
27.9
15.5
12.3

0.0
0.1
0.0
2.3
0.2
0.3
1.4
0.8
0.4
0.2
0.1
0.3
0.0
0.6
0.6
3.6
2.9
0.9
1.0
0.2
1.1
1.0

SFA, saturated fatty acids; UFA, unsaturated fatty acids; MUFA, monounsaturated fatty acids; PUFA, polyunsaturated fatty acids.

signicant roles in gonad maturation (i.e. gametogonesis and vitellogenesis).

Among 2530 fatty acid peaks, we were able to
securely identify 15 main fatty acids in pikeperch
liver. Unidentied fatty acids were of minor importance quantitatively and their ratios changed between
seasons from 14% to 3% (in male) and from 10% to
6% (in female). Palmitic acid is a major fatty acid
(Gibson et al. 1984) in sh and its level was not
inuenced by diet (Ackman et al. 1975). In the
present study, the relative proportions of SFA and
palmitic acid in liver were similar to those previously published results. Oleic acid (18:1) was the
most abundant monoenes in both sexes. High levels
of oleic acid and palmitoloic acid (16:1) have been
reported as a characteristic property of sh oils in
freshwater (Ackman 1967; Osman et al. 2001). In
this study, oleic acid level shows the characteristic
property of freshwater sh and palmitolic acid level
did not exceed the levels reported in marine sh. In
male liver, several fatty acids (C12:0, C16:0, C18:1,
C18:3n-3) were found greatly lower than their levels
in female liver in November. The decreases in SFA
(C12:0, C16:0) may be a consequence of the sharp
drops in temperatures, which, together with an
increase in PUFA, may be a physiological adaptation
to an extreme cold environment (Logue et al. 2000;
Hall et al. 2002; C
elik et al. 2005). On the other
hand, based on the previously published data
(Luzzana et al. 1996), it is assumed that oleic acid
(C18:1) and linolenic acid (C18:3n-3) may be
444

decreased on account of the preferential

utilisation for the testis development as determined
by GSI.
Substantial differences between the seasons were
observed in the relative proportion of PUFA proles
in the liver. Docosahexaenoic acid (22:6n-3) and
eicosapentaenoic acid (20:5n-3) were the most
abundant and varied n-3 PUFA in both sexes. In
parallel with several published studies indicating that
PUFA are abundant in sh from cold regions and
seasons (Dutta et al. 1985; Skuladottir et al. 1990;
Dey et al. 1993), the results presented here also
show that the levels of PUFA in male and female
sharply increased during the signicant decreases in
temperature in November (9.8 C). Although the
temperature continued to decrease in January
(6.4 C), the level of PUFA did not increase
accordingly, which was contrary to our expectations.
The decrease in the amount of PUFA in this period
may be because of the signicant utilisation of
PUFA and MUFA in gonad maturation, which may
overwhelm the stimulatory effects of temperature.
The inverse relationship between SFA and PUFA
observed in the dense feeding period may be
because of the upregulation of synthesis of PUFA
rather SFA.
Acknowledgements
The work was supported in part by a grant to K. Uysal from the
University of Suleyman Demirel, Isparta, Turkey.

Seasonal variations in pikeperch liver fatty acids

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