ISSN 1063-0740, Russian Journal of Marine Biology, 2008, Vol. 34, No. 1, pp. 51–56. © Pleiades Publishing, Ltd.

, 2008.
Original Russian Text © P.V. Velansky, E.Ya. Kostetsky, 2008, published in Biologiya Morya.

BIOCHEMISTRY

Lipids of Marine Cold-Water Fishes

P. V. Velansky and E. Ya. Kostetsky
Far Eastern State University, Vladivostok, 690000 Russia
e-mail: velansky@marbio.dvgu.ru
Accepted: September 13, 2007

Abstract—The study deals with the lipid and fatty acid compositions of the muscles, gills and liver of marine
fishes inhabiting cold waters (0.5–6°C) and caught in Peter the Great Bay (3 species) and Vostok Bay (2 spe-
cies), as well as with the fatty acid compositions of the main phospholipids in the muscle tissues of fish from
Olyutorskii Bay (4 species). The average content of phosphatidylcholine was about 60% in muscles and in the
liver and 53.8% of the sum of all phospholipids in gills. The phosphatidylethanolamine content was on the aver-
age 24.3, 25.1 and 22.3% in muscles, liver and gills, respectively. Increased contents of phosphatidylserine and
sphingomyeline were recorded in the gills. The mean (S.D.) molar ratios of cholesterol/phospholipids were
0.20, 0.32, and 0.58 in the muscles, liver, and gills, respectively. It was established that phosphatidylcholine has
a higher content of saturated fatty acids, whereas phosphatidylethanolamine was richer in monoenic acids and
docosahexaenoic acid (DHA). It was noted that the level of polyenic fatty acids was increasing and the level of
monoenic and saturated acids was decreasing in the series from gills–liver–muscles. The species with a more
active mode of life were distinguished by an increased content of docosahexaenoic acid.

Key words: phospholipids, fatty acids, cholesterol, phosphatidylcholine, phosphatidylethanolamine, docosa-

hexaenoic acid, cold-water fishes
DOI: 10.1134/S1063074008010070

Many papers have been published on the molecular
mechanisms of adaptation of fish cellular membranes
to various environmental factors. Most of the studies
have been performed either on fish during experimental
acclimation [11, 19], or in cell cultures [18]. These
studies concerned only a limited number of species and
gave a preference to fresh-water fishes [15, 20]. The
results thus obtained can hardly be compared with
results of investigations of fish in the natural environ-
ment. In this connection, our study is aimed at an anal-
ysis of the lipid composition of muscles, gills and liver
of different species of marine cold-water fish caught
directly in their natural habitat.
MATERIAL AND METHODS
The following fish species were used in the study:
the Pacific saffron cod Eleginus gracilis (Tilesius,
1810), the Korean flounder Glyptocephalus stelleri
(Schmidt, 1904) and the black-edged sculpin Gymno-
canthus herzensteini (Jordan et Starks, 1904) caught in
Peter the Great Bay at a water temperature of 3.4°C,
50 m deep; the black plaice Liopsetta obscura (Herzen-
stein, 1891) and the plain sculpin Myoxocephalus jaok
(Cuvier, 1829) caught in Vostok Bay, at a water temper-
ature of 0.5°C, 2 m deep. As well, in Olyutorsky Bay,
the Pacific cod Gadus macrocephalus (Tilesius, 1810)
(at a temperature of 4.2°C, 298 m deep), the Walleye
pollock Theragra chalcogramma (Pallas, 1811) (240 m
deep, at 4.2°C), the Atka mackerel, Pleurogrammus
monopterygius (Jordan et Metz, 1913) (167 m deep, at
6.1°C), and the Gray rockfish Sebastes glaucus (Hil-
dendorf, 1880) (142 m deep, at 4.3°C) were caught. We
chose no less than three specimens of every species and
combined their tissues. Immediately after the catch,
fish tissues were ground and fixed in a mixture of chlo-
roform–methanol (2 : 1 vol.). The total lipid extract was
obtained by the Folch method [8]. Qualitative analysis
of phospholipids (PL) was conducted with the use of
two-dimensional thin-layer chromatography (2-D
TLC) with the following quantification of phosphorus
in areas corresponding to individual PLs with the use of
a molybdate reagent for PLs [17]. The amount of cho-
lesterol was determined by the Liebermann–Burchard
reaction [3]. Phosphatidylcholine (PC) and phosphati-
dylethanolamine (PE) were isolated with the use of
one-dimensional TLC on silica gel in the chloro-
phormómethanol–water system (65 : 25 : 4, vol./vol.).
Fatty acids (FA) were analyzed as methyl esters. Meth-
ylation was conducted within 1 h at 90°C in anhydrous
solution of 5% HCl, purification of methyl ethers was
done by TLC on silica gel with a mobile phase consist-
ing of bensol. The analysis was carried out in an iso-
thermal mode on a 6890GC gas chromatograph with an
Innowax capillary column (30 m × 0.25 mm × 0.25 µm).
The temperature was programmed for 240°C on the
vaporizer, 200°C on the column, and 250°C on the
detector; the carrier gas was helium and the linear
velocity was 35 cm/sec. Identification was carried out
through calculation of the equivalent chain length) [6].

51
52 VELANSKY, KOSTETSKY
The contents of phospholipids and cholesterol were
determined based on the results of triplicate analyses,
the FA were quantified by the results of one chromato-
graphic examination.
The contents of phospholipids, cholesterol, and fatty
acid compositions of PC and PE were analyzed in the
muscles, gills and liver of fishes from the Sea of Japan.
In tissue samples from fishes caught in the Bering Sea,
only the fatty acid compositions of PC and PE were
studied.
RESULTS AND DISCUSSION
The content of PC in the organs and tissues studied
varied from 3.7 to 51.8% of the total lipids (Table 1).
The lowest content of PL was characteristic of the liver.
The amount of cholesterol also varied significantly:
from 0.6 to 13.8% of total lipids, with the lowest con-
tent of cholesterol being found in the liver (0.6–3.9%).
Despite a wide range of dispersion of the estimates for
cholesterol and phospholipids, their molar ratio varied
less distinctly and grew along the series: muscles
(0.16–0.21)–liver (0.2–0.34%)–gills (0.51–0.64%). A
high Chl/PL ratio in the gills was due to the fact that
Chl regulates the hydrophobic portion of the membrane
in order to reduce its osmotic permeability to water
[13]. According to literature data [2], the mean ratio of
cholesterol : phospholipids for the membranes of the
respiratory organs of teleosts is about 0.42.
The main phospholipids found were PC and PE,
they were seen in all the studied organs and tissues of
the fish. Other phospholipids found included sphingo-
myeline (SM), the hydrophobic portion of the mem-
brane phosphatidylserine (PS), phosphatidylinosite
(PI), diphosphatidylglycerol (DPG), lysophosphatidyl-
choline (LPC), and phosphatidic acid (PA). The mean
contents of PC in the muscle tissues and livers of the
fish were similar within the calculation error equaling
60.7 and 59.4%, respectively. For gills, this value was
somewhat lower, at 53.8%. The amount of SM, another
choline-containing lipid, grew along the series mus-
clesóliver–gills from 3.5 to 5.5 and 9.3%, respectively.
The sum of choline-containing lipids (PC, SM, and
LPC) in muscles, liver, and gills became closer at the
expense of SM to become 65.2, 65.4, and 64.0%. The
mean quantity of PE, with the calculation error consid-
ered, did not differ significantly and was 24.3, 25.1, and
22.3% in the muscles, liver, and gills, respectively. The
contents of PI and DPG in the muscles, liver, and gills
were similar: 5.6, 4.8, and 4.8%, respectively, for PI
and 1.2, 1.2, and 1.4% for DPG. We noted an enhanced
level of PS and SM in the gills.
An analysis of fatty acid composition of PC
(Table 2) showed that the main components of this PL
are palmitic (16 : 0), eicosapentaenoic (EPA, 20 : 5n-3),
and docosahexaenoic (DHA, 22 : 6n-3) acids. Their
total content in the gills, liver, and muscles equaled
41.9, 54.6, and 64.9%, on average respectively. The
RUSSIAN JOURNAL OF MARINE BIOLOGY
amount of the 16 : 0 acid varied insignificantly among
the organs.
The maximum contents of EPA and DHA were
recorded in muscles, and the minimal contents of these
acids were found in gills. The fatty acid composition of
PC showed a trend towards an increase in the mean
level of polyenic FAs along the series gillsóliver–mus-
cles (33.0, 42.3, and 52.6%, respectively) and towards
a decrease of the level of monoenoic (28.5, 21.2, and
15.9%), and saturated (37.0, 35.8, and 29.5%) FAs.
Our study of the fatty acid composition of PE
showed that the main components of the PL were 16 :
0, EPA, DHA, oleic (18 : 1n-9), and cis-vaccenic (18 :
1n-7) acids (Table 3). The pattern of distribution of FAs
among the organs is the same as that for PC: an
enhancement of the level of polyenic acids along the
series gillsóliver–muscles (39.9, 42.7, and 59.5%,
respectively) and an insignificant decrease in the level
of monoenic (31.3, 29.3, and 25.6%) and saturated
(27.9, 26.9, and 14.0%) acids along the same series.
The fatty acid composition of PE in the liver and gills
did not differ so distinctly as did PC.
In general, phosphatidylethanolamine is a more
unsaturated phospholipid, than phosphatidylcholine,
and the content of saturated fatty acids in phosphati-
dylethanolamine is 1.2–2.5 times lower. The level of
monoenic fatty acids in phosphatidylethanolamine is
1.1–2 times higher at the expense of the 18 : 1 acids
(especially cis-vaccenic acid) in all the tissues, while
phosphatidylcholine is richer in 16 : 1 monoenic fatty
acids. As compared to phosphatidylcholine, phosphati-
dylethanolamine contains higher contents of polyenoic
fatty acids in muscles (a mean of 59.5% versus 52.6%)
and in gills (39.9% versus 33.2%), while the estimate
for the liver was very similar, being approximately
42%. Higher contents of polyenoic fatty acids in the
muscles and in gills were found for different fatty acids.
So, compared with phosphatidylethanolamine, the
phosphatidylcholine of the muscles and liver contained
more EPA (19.7% versus 13.1% and 16.0% versus
12.4%, respectively); and the DHA content of all
organs was higher in phosphatidylethanolamine than in
phosphatidylcholine (34.9% versus 23.9% in muscles,
20.3% versus 15.6% in liver, and 13.7% versus 9.9% in
gills). These differences in the fatty acid compositions
of PC and PE were documented previously [10] and
seem to be characteristic of teleosts in particular [9].
This difference in the fatty acid compositions of PC
and PE in different organs should have some functional
meaning. It has been found that the PE of marine inver-
tebrates is more saturated; however, the phase transi-
tions of PC and PE lie in the same temperature range
[14]. At the same saturation level of fatty acid radicals,
the point of fusion for PE is higher by 20°C, than that
for PC, but fusion temperatures of PC and PE become
more similar at an increased unsaturation level [16].
Hence, considering that PE is usually localized in the
inner monolayer of the membrane, while PC is local-
Vol. 34 No. 1 2008
 LIPIDS OF MARINE COLD-WATER FISHES 53

Table 1. Contents of phospholipids and cholesterol in organs and tissues of five fish species from the Sea of Japan

Myoxocephalus Gymnocanthus Glyptocephalus Eleginus

Lipid Liopsetta obscura
jaok herzensteini stelleri gracilis

Muscles
PL 40.7 ± 4.8 51.8 ± 2.5 45.4 ± 2.1 28.3 ± 0.1 47.9 ± 0.6
Chl 4.4 ± 0.3 4.3 ± 0.8 5.0 ± 0.2 3.1 ± 0.1 4.9 ± 0.1
Chl/PL 0.21 ± 0.04 0.16 ± 0.04 0.21 ± 0.02 0.21 ± 0.01 0.20 ± 0.01
PC 58.3 ± 2.4 56.6 ± 2.4 64.1 ± 3.0 60.7 ± 0.7 63.9 ± 3.1
PE 24.6 ± 3.4 26.8 ± 1.2 24.6 ± 3.0 24.5 ± 0.3 21.1 ± 0.7
SM 3.7 ± 1.1 3.6 ± 0.5 3.4 ± 0.2 3.2 ± 0.7 3.9 ± 0.5
PS 4.5 ± 1.0 3.3 ± 0.5 2.1 ± 0.2 3.4 ± 0.5 2.9 ± 0.9
PI 5.7 ± 0.8 7.3 ± 0.8 2.8 ± 1.1 5.8 ± 0.7 6.2 ± 0.4
DPG 1.8 ± 0.4 1.2 ± 0.0 0.9 ± 0.2 1.0 ± 0.2 1.2 ± 0.3
LPC 0.8 ± 0.1 0.7 ± 0.2 2.0 ± 0.2 1.1 ± 0.2 0.6 ± 0.1
PA 0.6 ± 0.3 0.5 ± 0.2 0.1 ± 0.1 0.2 ± 0.1 0.2 ± 0.3
Liver
PL 6.0 ± 0.7 18.0 ± 1.3 19.0 ± 0.1 7.7 ± 0.1 3.7 ± 0.2
Chl 1.1 ± 0.0 3.9 ± 0.3 2.0 ± 0.2 1.2 ± 0.1 0.6 ± 0.1
Chl/PL 0.34 ± 0.05 0.42 ± 0.07 0.20 ± 0.03 0.31 ± 0.04 0.32 ± 0.07
PC 58.0 ± 2.91 59.4 ± 1.8 62.3 ± 1.5 57.3 ± 1.3 60.1 ± 2.8
PE 21.1 ± 1.6 26.9 ± 1.2 26.0 ± 0.3 22.8 ± 1.4 28.4 ± 2.2
SM 6.4 ± 0.3 5.2 ± 0.5 4.3 ± 0.2 6.0 ± 0.5 5.7 ± 0.5
PS 4.5 ± 0.2 4.3 ± 1.0 2.0 ± 2.1 3.8 ± 0.5 1.8 ± 0.2
PI 7.4 ± 0.9 2.5 ± 0.2 4.5 ± 3.2 8.0 ± 1.3 1.7 ± 0.4
DPG 1.3 ± 0.2 1.1 ± 0.1 0.7 ± 0.3 1.5 ± 0.1 1.5 ± 0.0
LPC 1.0 ± 0.7 0.3 ± 0.2 0.2 ± 0.2 0.3 ± 0.1 0.6 ± 0.1
PA 0.3 ± 0.1 0.3 ± 0.2 0.0 ± 0.0 0.3 ± 0.1 0.3 ± 0.2
Gills
PL 36.8 ± 1.4 45.6 ± 5.7 15.6 ± 1.5 23.1 ± 1.4 23.3 ± 0.6
Chl 9.7 ± 1.1 13.8 ± 1.7 5.2 ± 0.1 6.8 ± 0.1 6.8 ± 0.5
Chl/PL 0.51 ± 0.08 0.59 ± 0.13 0.64 ± 0.08 0.57 ± 0.04 0.56 ± 0.06
PC 51.1 ± 2.0 53.8 ± 1.9 53.9 ± 3.0 55.3 ± 1.9 55.0 ± 2.8
PE 24.9 ± 2.8 23.3 ± 1.8 22.7 ± 2.0 18.5 ± 1.1 21.9 ± 2.1
SM 8.1 ± 0.3 11.2 ± 0.9 9.0 ± 0.6 9.8 ± 1.0 8.6 ± 2.0
PS 8.0 ± 0.8 5.1 ± 1.3 7.0 ± 1.3 6.3 ± 0.7 6.5 ± 0.6
PI 5.5 ± 0.7 3.6 ± 0.7 4.2 ± 0.5 5.1 ± 0.1 5.7 ± 0.9
DPG 1.4 ± 0.4 1.7 ± 0.4 1.3 ± 0.1 1.5 ± 0.0 1.3 ± 0.1
LPC 0.3 ± 0.1 1.2 ± 1.0 1.5 ± 0.6 1.5 ± 0.4 0.2 ± 0.1
PA 0.7 ± 0.3 0.1 ± 0.1 0.4 ± 0.6 1.9 ± 0.6 0.8 ± 0.4
Note: PL and Chl in % of the total lipid content; individual PLs in % of the total PL content; Chl/PL presented as a molar ratio.

RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 34 No. 1 2008

54 VELANSKY, KOSTETSKY

Table 2. Fatty acid composition of PC in muscles, liver, and gills (% of the total fatty acid content)
Fatty Acid

monoenic
16 : 1 n-9

16 : 1 n-7

18 : 1 n-9

18 : 1 n-7

18 : 2 n-6

20 : 1 n-9

20 : 4 n-6

20 : 5 n-3

22 : 5 n-3

22 : 6 n-3

saturated

polyenic
Species

14 : 0

16 : 0

18 : 0
Muscles
Gadus macrocephalus 1.4 23.5 1.2 2.0 7.7 9.1 4.3 1.8 0.3 2.7 12.8 1.4 21.7 34.3 19.2 42.4
Theragra chalcogramma 1.2 19.7 0.4 1.5 5.1 7.8 2.9 0.9 0.6 1.7 16.6 1.7 31.8 26.8 15.5 54.6
Pleurogrammus monop- 0.9 15.7 0.5 1.8 8.9 6.2 2.2 1.1 1.0 1.7 17.3 1.5 31.5 27.3 14.2 54.9
terygius
Sebastes glaucus 0.9 19.4 0.3 1.6 8.0 5.3 1.9 1.4 0.2 2.1 14.7 1.3 35.7 29.5 11.4 56.8
Myoxocephalus jaok 0.6 18.3 1.2 3.1 3.1 8.6 5.6 0.6 0.7 5.1 22.3 3.3 19.1 24.1 21.9 53.8
Liopsetta obscura 0.7 28.2 1.7 1.4 4.1 3.5 2.0 0.3 0.3 5.6 23.9 3.2 16.1 35.2 12.0 52.6
Gymnocanthus herzen- 0.9 22.0 0.6 3.0 2.6 5.8 3.6 0.4 0.3 2.9 28.3 3.0 15.8 27.7 17.1 54.4
steini
Glyptocephalus stelleri 1.7 21.8 0.6 1.9 4.2 7.7 3.2 0.5 0.7 3.6 22.2 2.9 17.4 30.7 17.0 50.5
Eleginus gracilis 0.9 23.0 0.6 2.3 3.9 6.1 3.4 0.4 0.4 2.7 19.1 1.7 25.9 29.9 15.9 53.5
Liver
Myoxocephalus jaok 1.8 20.6 2.3 4.8 4.2 8.4 4.0 1.1 0.6 3.4 14.5 3.0 16.7 31.8 24.0 43.6
Liopsetta obscura 1.9 29.4 2.9 2.2 6.9 3.7 2.1 0.8 0.3 3.6 14.7 2.5 14.4 44.3 14.4 40.3
Gymnocanthus herzen- 1.7 18.2 1.4 4.9 4.7 6.9 4.0 0.3 0.2 3.3 24.2 4.2 14.3 28.4 20.9 50.2
steini
Glyptocephalus stelleri 3.6 24.0 2.2 3.2 5.2
8.2 3.7 1.5 0.7 2.1 13.0 2.4 13.1 39.7 22.9 36.4
Eleginus gracilis 2.0 23.2 1.3 4.1 4.5
8.9 3.9 0.6 1.0 2.5 13.4 1.0 19.4 34.6 24.1 40.8
Gills
Myoxocephalus jaok 0.8 15.6 1.4 7.9 6.2 10.1 7.1 0.6 1.3 8.0 15.0 2.7 12.9 24.9 31.5 43.0
Liopsetta obscura 2.3 19.6 3.4 3.8 10.8 7.4 2.4 0.1 0.8 5.7 9.9 2.4 11.6 41.2 22.0 35.3
Gymnocanthus herzen- 2.4 23.6 2.1 8.3 7.2 12.0 6.2 0.7 1.1 2.8 10.8 1.4 4.9 39.0 36.2 23.9
steini
Glyptocephalus stelleri 3.5 22.7 3.3 3.5 6.6 11.5 4.9 1.7 1.1 3.3 10.1 2.2 7.8 40.2 29.7 28.2
Eleginus gracilis 1.8 20.9 1.6 2.4 5.8 8.8 3.2 0.8 1.3 2.9 11.8 1.1 12.5 39.9 23.2 35.8
Note: Fatty acids with content below 1% in all the samples are not presented in the table.

ized in the outer membrane layer [12], we can suggest
that higher PE unsaturation compensates for the differ-
ences in the phase transition temperatures of the two
main phospholipids. This, in its turn, is responsible for
the phase of the liquid-crystalline state of the outer and
inner layers of the membrane and, consequently, for the
synchronism of membrane function.
Interspecies differences in FA composition allowed
us to distinguish species having high DHA levels: Ther-
agra chalcogramma, Pleurogrammus monopterygius,
Sebastes glaucus, and Eleginus gracilis. It is known
that DHA does not participate in thermoadaptation [2,
5] and that the DHA level depends on the intensity of
metabolism [4], i.e., directly on the motility of the fish.
Our data support this supposition. Among the species
studied, we found the maximum DHA contents in ben-
thopelagic fishes with an active life mode: T. chalco-
gramma, P. monopterygius, and E. gracilis. The
benthic species S. glaucus was an exception: its muscle
tissue contained the maximum amount of DHA and
polyenoic FA among all other species.
It is accepted that the main fatty acids participating
in the thermoadaptation of fish are 18 : 1 [7] and EPA
[2]. The fish species studied inhabit waters of low tem-
peratures from 0.5 to 6.1°C. It is evident therefore, why
the contents of 18 : 1 in PC and EPA in PC and PE do
not vary considerably within one tissue among the spe-
cies. The only exception was Liopsetta obscura that
had a relatively low level of 18 : 1 in the PC. Markedly
enhanced amounts of EPA were found in the liver and
muscles of Gymnocanthus herzensteini and in the gills
of Myoxocephalus jaok, with the cholesterol : phospho-
lipids ratio in these samples being lower than in other
species.

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 LIPIDS OF MARINE COLD-WATER FISHES 55

Table 3. Fatty acid composition of PE in muscles, liver, and gills (% of the total fatty acid content)
Fatty Acid

monoenic
16 : 1 n-9

16 : 1 n-7

18 : 1 n-9

18 : 1 n-7

18 : 2 n-6

20 : 1 n-9

20 : 4 n-6

20 : 5 n-3

22 : 5 n-3

22 : 6 n-3

saturated

polyenic
Species

14 : 0

16 : 0

18 : 0
Muscles
Gadus macrocephalus 0.7 11.6 0.3 0.9 4.3 8.4 6.8 1.3 1.0 1.7 11.9 2.0 38.4 17.9 20.0 57.7
Theragra chalcogramma 0.6 10.5 0.3 0.8 3.3 8.4 5.5 1.0 1.9 1.0 12.3 2.2 42.2 15.4 21.0 60.9
Pleurogrammus monop- 0.9 11.0 0.3 1.2 5.3 7.0 3.7 1.5 3.5 1.4 11.7 1.9 36.1 19.2 21.5 55.5
terygius
Sebastes glaucus 0.5 10.8 0.2 0.6 4.8 6.5 2.7 3.3 0.6 1.0 7.6 1.2 47.9 18.1 14.3 64.2
Myoxocephalus jaok 0.3 5.8 0.5 2.1 4.4 11.4 12.3 1.3 2.2 5.0 13.3 4.1 26.9 12.7 31.6 55.2
Liopsetta obscura 0.4 9.3 0.6 1.4 4.9 8.3 7.1 1.1 1.2 4.4 14.5 4.9 28.5 16.8 24.3 58.4
Gymnocanthus herzen- 0.2 4.5 0.2 1.4 4.3 6.1 10.4 0.5 1.1 3.4 20.1 4.7 28.1 11.2 24.1 63.2
steini
Glyptocephalus stelleri 0.3 5.2 0.2 1.7 4.7 10.6 12.8 0.7 2.3 5.5 13.6 4.0 25.7 12.9 31.4 54.4
Eleginus gracilis 0.4 8.4 0.2 0.7 5.1 5.2 5.9 0.4 1.0 2.0 14.6 3.1 40.6 16.3 16.7 66.4
Liver
Myoxocephalus jaok 0.8 10.1 1.1 3.3 4.4 13.8 12.9 1.0 2.1 3.5 12.6 3.2 17.5 20.7 37.4 41.0
Liopsetta obscura 1.8 18.6 1.6 1.6 6.2 5.4 5.7 1.3 1.2 2.8 8.9 1.5 20.2 36.9 21.0 39.8
Gymnocanthus herzen- 0.4 8.2 0.3 2.4 4.2 8.4 14.2 0.4 0.7 1.7 19.5 4.1 21.2 18.4 31.0 50.4
steini
Glyptocephalus stelleri 2.2 15.4 1.9 3.5 7.5 11.2 6.0 1.5 1.7 1.9 10.8 2.5 16.4 31.4 30.2 37.2
Eleginus gracilis 0.9 11.3 0.7 1.5 6.5 8.0 8.3 1.2 2.4 1.5 10.3 1.6 26.4 27.0 26.7 45.3
Gills
Myoxocephalus jaok 0.5 5.4 0.7 2.9 4.1 8.2 13.3 0.7 3.5 9.1 16.8 3.2 18.4 12.8 33.3 53.6
Liopsetta obscura 1.3 13.7 0.8 2.4 9.0 11.1 6.3 1.4 2.7 6.7 11.2 2.8 15.8 28.2 28.3 42.6
Gymnocanthus herzen- 1.8 15.5 1.0 3.5 8.2 9.0 14.3 1.2 2.5 3.7 10.5 2.1 6.8 32.9 37.5 28.8
steini
Glyptocephalus stelleri 1.1 19.9 2.1 2.1 8.0 15.2 4.9 5.6 2.5 3.6 7.4 2.4 9.2 35.7 32.5 31.1
Eleginus gracilis 1.3 12.1 1.3 2.0 5.7 7.4 6.4 0.6 2.6 3.5 11.0 2.0 18.3 29.7 25.1 43.5
Note: Fatty acids with content below 1% in all the samples are not presented in the table.

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