Metoyo Et Al, 2005

Effect of dietary fish oil substitution with linseed oil on the performance, tissue
fatty acid profile, metabolism, and oxidative stability of Atlantic salmon1,2

D. Menoyo,* C. J. Lopez-Bote,3 A. Obach, and J. M. Bautista*
Departamento de *Bioqumica y Biologia Molecular IV and Produccion Animal,
Facultad de Veterinaria, Universidad Complutense de Madrid, 28040 Madrid, Spain; and
Nutreco Aquaculture Research Centre, N-4001 Stavanger, Norway
ABSTRACT: The objective of this experiment was to

test the effect of total or partial substitution of dietary
fish oil (FO) by linseed oil (LO) in Atlantic salmon feeding on performance, liver and muscle fatty acid composition, selected lipogenic and lipolytic enzyme activities,
and flesh oxidative stability. For 12 wk, fish (220 12
g of initial BW) were fed five experimental diets in
which the FO was serially replaced by 25, 50, 75, and
100% LO. Total FO replacement by LO did not (P =
0.20) affect fish final weight, biometric indices, or i.m.
fat contents. Liver and muscle neutral lipid (NL) composition responded to dietary treatments in different
ways. Whereas the sum of n-3 PUFA in muscle followed
a linear and quadratic pattern with increasing levels
of LO, a linear (P = 0.005) effect was observed in the
liver NL fraction. Total n-3 and n-6 PUFA contents in
the polar lipid fraction (PL) were unaffected (P = 0.356)

by dietary input of LO in muscle. Activity of liver glucose-6-P-dehydrogenase (G6PD) was greater with increasing levels of LO (P = 0.004). A time effect (P <
0.001) was observed in the concentration of lipid peroxidation products, expressed as thiobarbituric acid reactive substances, in fish flesh stored under refrigeration
for 9 d; however, the progressive inclusion of LO in the
feed did not affect (P = 0.125) flesh oxidation stability. In
summary, LO can totally replace FO in Atlantic salmon
feed without affecting growth performance and muscle
susceptibility to lipid oxidation. Fatty acid metabolism
in the liver was affected by LO, promoting G6PD activity and eicosatetraenoic acid accumulation; however, a
100% LO replacement decreased (P < 0.001) concentrations of eicosapentaenoic and docosahexaenoic acids in
salmon muscle.
Key Words: Atlantic Salmon, Fatty Acid, Lipid Metabolism, Vegetable Oil
2005 American Society of Animal Science. All rights reserved.
Introduction
Fish oil (FO) is a common fat source in fish diets
because of its high proportion of long-chain, n-3 fatty
acids, which are nutritionally essential to teleosts
(NRC, 1993). Because of predictable insufficient FO
availability for fish feeding, however, alternative
sources must be assessed (FAO, 2002). Studies performed in Atlantic salmon fed lipid rich diets containing
high-oleic sunflower oil (Torstensen et al., 2000), rapeseed oil (Bell et al., 2001), palm oil (Bell et al., 2002),
soybean oil (Grisdale Helland et al., 2002), and linseed
oil (LO; Bell et al., 2003b) were found to have no detri-
1
This research was partially financed by Ministerio de Ciencia y
Tecnologa (CICYT AGL-2001-1162).
2
The authors are grateful to R. Prieto for technical assistance.
3
Correspondencephone: 34-91 394 3889; fax: 34-91-394 3824;
e-mail: clemente@vet.ucm.es.
Received November 4, 2004.
Accepted August 1, 2005.
J. Anim. Sci. 2005. 83:28532862
mental effects on growth. Nonetheless, use of vegetable

oil sources in fish feeding still needs to be evaluated
in terms of flesh quality and metabolic use, especially
whether the fish has the ability to elongate linoleic
acid (18:3 n-3) to long-chain, n-3 fatty acids. Bell et al.
(2003b) reported a significant loss of n-3 highly unsaturated fatty acids (HUFA) in the flesh of Atlantic salmon
when FO was replaced with >66% of vegetable oils.
Conflicting reports on the effects of vegetable oils on
lipogenic and lipolytic enzymes (Torstensen et al., 2000;
Regost et al., 2003; Menoyo et al., 2003) suggest that
different vegetable oils may have different effects on
fish metabolism and that different species respond in
different ways. In fish, the pentose phosphate pathway
is active in the liver, where it provides the cytoplasmic
reducing equivalents (NADPH; Alvarez et al., 2000). In
the same way, fatty acid -oxidation is active in fish
livers, where it is regulated by mitochondrial uptake of
long-chain fatty acids through the activity of carnitine
palmitoyltransferase I (Fryland et al., 1998). Therefore, the objectives of this research were 1) to evaluate
the influence of dietary fat source (FO vs. LO) and com-
2853
2854
Menoyo et al.
Table 1. Composition of experimental diets

Linseed oil, % replacement of fish oil
Ingredient, g/kg of DM
Fish meal-LT (71.8/8.4)
Corn gluten
Wheat
Fish oil
Linseed oil
Mineral premixb
Vitamin premixb
Carophyll pink
-Tocopherolc
-Tocopherolc
25
50
75
100
632.0
30.1
70.0
259.4
5.6
2.2
0.6
321.4
9.6
632.0
30.1
70.0
194.6
64.9
5.6
2.2
0.6
323.8
30.3
632.0
30.1
70.0
129.7
129.7
5.6
2.2
0.6
318.9
41.6
632.0
30.1
70.0
64.9
194.6
5.6
2.2
0.6
318.6
50.2
632.0
30.1
70.0
259.4
5.6
2.2
0.6
350.1
80.2
LT = low temperature (71.8 CP/8.4 moisture)

Active ingredients supplied per kilogram of feed. Minerals: 48 mg of CuSO4, 2.0 mg of KI, 109.4 mg of
MnSO4, 0.4 mg of Na2SeO3, and 257.1 mg of ZnSO4 (all mineral salts from Sigma, St. Louis, MO). Vitamins:
5 mg of retinol acetate, 4.8 mg of vitamin D3, 300 mg of -tocopheryl acetate, 15.3 mg of thiamin-Cl, 26 mg
of riboflavin, 15.3 mg of pyridoxine-Cl, 88.9 mg of Ca-pantothenate, 153.1 mg of niacin, 5.2 mg of folic acid,
20.0 mg of vitamin B12, 150.0 mg of biotin, 408.2 mg of myo-inositol, 40.0 mg of vitamin K3, and 1 g of
ascorbate polyphosphate (all vitamins from Hoffmann-La Roche, Basel, Switzerland).
c
Analyzed concentration, mg/kg of DM.
b
binations for Atlantic salmon on fatty acid composition

and liver metabolism, and 2) to relate these findings to
susceptibility of flesh to lipid peroxidation.
Materials and Methods

Fish Husbandry and Feeding
The trial was carried out at the Nutreco Aquaculture
Research Center Lerang Research Station, Jrpeland,
Norway. A total of 110 Atlantic salmon (Salmo salar)
AquaGen strain, weighing approximately 220 g, were
distributed randomly into five 1-m 1-m circular tanks
containing 500 L of sea water and were fed one of the
five experimental diets for 12 wk. Fish were fed daily
to satiation, and feed disappearance was monitored
throughout the trial. Wasted feed was collected from
the effluent water from each tank by a wire mesh collector and dried. Net feed intake was registered daily. Fish
were subjected to a photoperiod regimen of 18 h light
and 6 h dark, and the temperature over the whole experimental period ranged from 8 to 9C. Experimental
diets were produced at the Nutreco Technology Center
(Stavanger, Norway) as extruded, sinking, 4-mm pellets. Basal composition of diets was the same except
for the oil added during vacuum fat coating (Table 1).
Batches of extruded pellets were produced from a common meal mixture and coated with FO that was serially
replaced by 25, 50, 75, or 100% LO. The fatty acid
composition of the experimental diets is presented in
Table 2, and diets were formulated to contain targeted
levels of 37% CP and 36% crude fat (DM basis).
Biological Parameters and Sample Collection

During the final sampling, 11 fish per tank were
killed by a blow to the head and immediately exsanguinated in chilled seawater; weight and fork length were
recorded. Then, fish were eviscerated, and the weight

of viscera and liver was registered to assess the hepatoand viscero-somatic indexes. Livers were then cut into
two pieces, and one portion was placed in liquid N2 and
stored at 80C for enzymatic analyses, whereas the
other half was frozen and stored at 0C for fatty acid
analysis. Fish were filleted, and left fillets were immediately vacuum-packaged and frozen at 20C for fatty
acid analysis and lipid peroxidation tests.
Chemical Analyses of Flesh Samples and Lipid

Peroxidation Assessment
Fatty acids of diets were extracted and quantified
by the one-step procedure described by Sukhija and
Palmquist (1988) in freeze-dried samples with pentadecanoic acid (15:0; Sigma, Alcobendas, Madrid, Spain)
as internal standard. Neutral lipids (NL) and polar
lipids (PL) from individual fillet and liver samples (five
fish per tank) were extracted using the method of
Marmer and Maxwell (1981). Before the analysis of
fatty acids by gas chromatography, all lipid samples
were methylated as described by Lopez-Bote et al.
(1997). Fatty acid methyl esters were then analyzed
using gas chromatography (Model HP-6890; Hewlett
Packard Co., Avondale, PA) equipped with flame ionization detection and a 30-m 0.32-mm 0.25-mm crosslinked polyethylene glycol capillary column (HewlettPackard-Innowax). Results were expressed as the percentage of each fatty acid with respect to the total
fatty acids.
Lipid peroxidation products were determined as thiobarbituric acid reactive substances (TBARS) in fish
flesh according to the procedure of Menoyo et al. (2002),
and TBARS were expressed as mol of malonaldehyde
(MDA)/kg of wet tissue. Vitamin E isoforms were extracted and quantified in feed and muscle as described
by Rey et al. (2001).
2855
Linseed oil in salmon feeding
Table 2. Fatty acid composition (%) of experimental diets

Fatty acid
Myristic acid (14:0)
Palmitic acid (16:0)
Stearic acid (18:0)
Total SFA
Palmitoleic acid (16:1n-7)
Oleic acid (18:1n-9)
Vaccenic acid (18:1n-7)
Cetoleic acid (22:1n-11)
Total MUFA
Linoleic acid (18:2n-6)
Arachidonic acid (20:4n-6)
Total n-6 fatty acids
Linolenic acid (18:3n-3)
Stearidonic acid (18:4n-3)
Eicosatrienoic acid (20:3n-3)
Eicosatetraenoic acid (20:4n-3)
Eicosapentaenoic acid (20:5n-3)
Docosapentaenoic acid (22:5n-3)
Docosahexaenoic acid (22:6n-3)
n-3:n-6 ratio
n-3 HUFAa
UIb
25
50
75
100
6.9
19.8
4.0
31.7
7.6
10.5
3.1
2.6
27.5
3.6
0.9
4.9
1.2
2.9
0.1
0.9
13.6
1.6
15.3
35.7
14.4
7.3
31.5
2.27
4.9
15.6
3.7
24.9
5.4
15.2
2.7
2.3
28.8
6.9
0.7
7.8
13.1
2.1
0.1
0.6
9.7
1.2
11.5
38.4
18.6
4.9
23.1
2.20
3.8
13.6
3.8
21.7
4.3
14.2
2.2
2.1
25.6
8.5
0.5
9.2
23.0
1.6
0.1
0.5
7.7
0.9
9.5
43.4
17.2
4.7
18.7
2.24
2.7
11.6
3.7
18.4
3.1
15.5
1.9
2.1
25.2
10.1
0.4
10.6
30.1
1.2
0.1
0.3
5.7
0.7
7.5
45.6
18.3
4.3
14.3
2.21
1.1
8.7
3.7
13.5
1.3
17.5
1.4
1.7
23.9
12.7
0.2
13.0
41.6
0.5
0.1
0.2
2.6
0.3
4.3
49.5
19.6
3.8
7.4
2.18
Highly unsaturated fatty acids (HUFA) include 20:3, 20:4, 20:5, 22:5, and 22:6.
UI = unsaturation index (average number of double bonds per fatty acid residue).
Mitochondrial Preparations, Soluble Extracts,

and Enzyme Analyses
Mitochondrial extracts from liver were prepared as
described by Menoyo et al. (2004), and the activity of
carnitine palmitoyltransferase I (CPT I) was assayed
according to procedures of Sanz et al. (2000). Liver homogenates and the activity of glucose-6-phosphate dehydrogenase (G6PD) were assayed as described by
Bautista et al. (1988). All enzyme activity assays were
performed in duplicate or triplicate at 30C. Enzymatic
activity units (IU), defined as moles of substrate converted to product per minute at the assay temperature,
were expressed per milligram of hepatic soluble protein
(specific activity). Soluble protein content of liver homogenates was determined by the method of Bradford
(1976), using BSA as the standard. The L [methyl3
H] carnitine hydrochloride (82.0 Ci/mmol) used in the
CPT I determination was supplied by Amersham Pharmacia Biotech (Barcelona, Spain), whereas Sigma supplied the remaining reagents.
Statistical Analyses
Data were analyzed as a completely randomized design by the GLM procedure of SAS (SAS Inst., Inc.,
Cary, NC), with LO inclusion level as the lone fixed
effect, and individual fish as the experimental unit. The
Tukeys test was used to separate treatment means,
and regression analysis was used to measure the linear
(L) or quadratic (Q) response to LO inclusion level.
A repeated measure mean test was used to compare

differences in TBARS concentrations between groups
during stimulated lipid peroxidation (Morris, 1999).
Results and Discussion

Growth performance and dorsal muscle composition
are presented in Table 3. No (P 0.20) significant effect
of dietary treatment was observed for final weight, condition factor, or viscero- and hepato-somatic indexes.
This is in accordance with previous reports showing
that feeding Atlantic salmon current commercial diets
with 45 to 50% CP (provided mainly as fish meal) vegetable oils can replace portions of the FO without negative effects on growth and biometric indexes (Torstensen et al., 2000; Grisdale Helland et al., 2002; Bell et
al., 2003b). Although some concern exists that feeding
diets with vegetable oil may affect fish health (Bell et
al., 1991; Thompson et al., 1996), no mortalities were
recorded in the current study. Neutral (mainly triglycerides) and polar (mainly phospholipids) i.m. lipids did
not (P > 0.07) differ among experimental groups. Triglycerides are the predominant lipid class in salmon
muscle (Bell et al., 2003a). Results of this study suggest
that LO does not promote changes in flesh adiposity
compared with FO, thereby differing from results observed in fish of the same size with palm oil and rapeseed oil (Bell et al., 2001, 2002). Vegetable oils affect
Atlantic salmon muscle fat content in different ways,
with palm oil and 50% rapeseed oil decreasing muscle
2856
Menoyo et al.
Table 3. Effect of dietary fish and linseed oil combinations on weight, biometry measurements, and muscle and liver neutral lipids (NL) and polar lipids (PL)
Item
Weight, g
Length, cm
Condition factor, %
VSI, %b
HSI, %c
Muscle NL, %
Muscle PL, %
Liver NL, %
Liver PL, %
25
50
75
100
Pooled SEa
P<F
525.86
33.28
1.43
11.49
1.50
5.87
1.25
3.07x
1.26
543.00
33.66
1.42
11.47
1.42
7.34
1.19
3.53xyz
1.38
526.81
33.19
1.44
11.31
1.48
6.24
0.92
3.44xy
1.46
537.71
33.60
1.42
10.96
1.45
5.61
0.90
5.48z
1.31
556.07
33.52
1.47
11.31
1.45
7.08
0.95
5.31yz
1.31
13.54
0.27
0.02
0.17
0.03
0.43
0.15
0.47
0.06
0.396
0.376
0.205
0.488
0.723
0.071
0.300
0.003
0.149
a
n = 11 for weight, length, condition factor, VSI, and HSI; n = 5 for muscle NL, muscle PL, liver NL, and
liver PL.
b
Viscero-somatic index (VSI) = carcass weight BW1 100.
c
Hepato-somatic index (HSI) = liver weight BW1 100.
x,y,z
Means within a row with different superscripts differ, P < 0.05.
adiposity (Bell et al., 2001, 2002), whereas LO had no

effect on muscle adiposity. It is interesting to note that
intrahepatic NL concentrations were greater (P < 0.003)
in groups receiving either 75 or 100% LO than in those
receiving 100% FO. This adiposity effect of vegetable
oil in salmon liver also was observed when feeding a
blend of LO and rapeseed oil (Tocher et al., 2001) or
when rapeseed replaced >50% of FO (Bell et al., 2001);
however, the same effect was not observed with palm
oil (Bell et al., 2002).
Liver G6PD and CPT I activities are shown in Figure
1. The activity of G6PD, which is the main donor of
reducing power in the form of NADPH, increased (P =
0.004) with increasing levels of LO, showing a specific
activity of 0.09 IU/mg of soluble protein in the liver of
fish fed 0% LO and reaching the maximum activity of
0.15 IU/mg of soluble protein in fish fed diets with 100%
LO. It has been well documented that the increasing
elongation activity in Atlantic salmon is triggered by
LO (Bell et al., 1993; Tocher et al., 2000). According to
this, we observed the accumulation of eicosatetraenoic
acid (20:4 n-3), which is the main product from the 6
desaturation of linolenic acid (18:3 n-3) in the liver
NL and PL fractions (Tables 4 and 5), with increasing
inputs of LO. Thus, the increase in G6PD activity may
be related to the greater concentration of intracellular
NADPH needed to accomplish the elongation process.
The carnitine shuttle, mediated by CPT I, is needed for
long-chain fatty acids to cross the inner mitochondrial
membrane and is tightly controlled by its inhibition of
elevated levels of cellular malonyl-CoA (Zammit, 1999),
the two-carbon donor needed for the elongation process.
It is plausible, therefore, to relate fatty livers to a
greater elongation process; however, we were unable
to detect differences on CPT I activity in the liver (Figure 1).
It is generally accepted that dietary fatty acid profile
is closely reflected in fatty acid composition of fish tissues (Rosenlund et al., 2001a; Bell et al., 2002; Caballero et al., 2002). Linseed oil (Table 2) provided a lower
concentration of total SFA, and a higher concentration

of total n-6 fatty acids and total n-3 fatty acids than
FO. As expected, n-3 fatty acids provided by LO were
of shorter chain length (41.6% 18:3 n-3) and had less
unsaturation than n-3 fatty acids provided by FO
(13.6% eicosapentaenoic acid [20:5 n-3] and 15.3% docosahexaenoic acid [22:6 n-3]). It is interesting to note,
Figure 1. Effect of increasing linseed oil (% replacement

of fish oil) in the feed of Atlantic salmon on liver glucose6-phosphate dehydrogenase (G6PD) and carnitine palmitoyltransferase I (CPT I) activities. Bars that do not have
a common letter differ, P = 0.004.
2857
Table 4. Effect of dietary fish and linseed oil combinations on selected fatty acids (g/100
g of total fatty acids) of neutral hepatic lipids
Item
No. of fish
Stearic acid (18:0)
Total SFA
Total MUFA
Homo--linolenic acid (20:3n-6)
n-3:n-6 ratio
n-3 HUFAa
UIb
25
50
75
100
Pooled SE
P<F
5
1.84z
14.95z
7.02z
24.29z
2.77z
10.40x
0.42
19.48x
2.13w
0.26z
3.94z
6.34x
0.84w
0.39
0.27x
1.59xy
10.57z
4.27z
31.31z
49.24yz
13.42y
7.79z
48.01z
3.15z
5
1.64z
13.62z
6.66yz
22.30z
2.25yz
13.33x
0.57
21.68xy
3.67x
0.21y
2.96y
6.85x
4.27x
0.37
1.35xy
1.26x
7.88y
3.34y
29.65z
48.13xyz
17.17y
7.07z
43.49z
3.07z
5
1.30z
13.93z
6.53yz
22.08z
1.83xy
11.66x
0.42
18.55x
3.82x
0.17x
3.08y
7.07x
6.99x
0.32
2.25y
1.42xy
7.86y
2.79x
29.54z
51.17z
15.19y
7.24z
43.86z
2.98z
5
1.32z
10.05y
5.66xy
17.30y
2.10yz
19.01y
0.80
27.82yz
6.70y
0.18x
1.65x
8.53y
13.96y
0.39
4.01z
1.69y
4.91x
1.94w
17.89y
44.78xy
24.00z
5.28y
30.44y
2.56y
5
0.66y
7.41x
5.26x
13.45x
1.31x
24.61z
0.50
31.69z
8.50z
0.10w
0.89x
9.50z
18.21z
0.54
4.72z
2.58z
3.42w
1.00v
13.09y
43.57x
29.87z
4.59y
24.82y
2.39y
0.14
0.64
0.23
0.75
0.19
1.40
0.12
1.85
0.36
0.04
0.22
0.19
0.76
0.07
0.30
0.09
0.42
0.13
1.84
1.51
1.67
0.31
2.06
0.09
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
0.407
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
0.125
<0.001
<0.001
<0.001
<0.001
<0.001
0.002
<0.001
<0.001
<0.001
<0.001
Within a row, means that do not have a common superscript letter differ, P < 0.05.
v,w,x,y,z
however, that concentrations of myristic (14:0) and palmitic (16:0) acids were considerably greater in FO than
in LO (6.9 and 19.8% vs. 1.1 and 8.7%, respectively),
thereby leading to a general overall unsaturation (as
assessed by the unsaturation index) and average fatty
acid chain length similar between fat sources (2.27 vs.
2.18 and 18.3 vs. 18.1 for FO and LO, respectively).
Diets with combinations of LO and FO showed intermediate values, and, interestingly, the unsaturation index
decreased (P < 0.05) in the i.m. NL fraction when LO
replaced FO (Table 4).
Hepatic and i.m. NL and PL fatty acid compositions
are shown in Tables 4, 5, 6, and 7, respectively. The
amount of MUFA was greater (P < 0.05) in the i.m. NL
fraction than in the diet in response to the accumulation
of oleic acid (18:1n-9). Selection of a given fatty acid to
provide energy will depend on availability related to
the amount and interactions with other dietary fatty
acids (Bell et al., 2002). In general terms, when provided
in excess, MUFA and 16:0 seem to be readily used for oxidation in salmonids muscles. Similarly, linoleic acid
(18:2 n-6), 18:3 n-3, and 20:5 n-3 are well oxidized when
high amounts are included in the diet, which may be
caused by a surplus of essential fatty acids (Kiessling
and Keissling, 1993; Mckenzie et al., 1998; Bell et al.,
2003b). Although oxidation could explain the relative
high accumulation of 18:1n-9 in muscle samples in the
current study, other metabolic regulation processes,
such as changes in 9 desaturase activity, also might

be involved; however, accumulation of 18:1n-9 was different in the NL fraction of liver and muscle (Tables 4
and 5). Whereas it followed a linear (R2 = 0.91; P <
0.001) pattern in muscle, a linear (R2 = 0.73; P < 0.001)
and quadratic effect (R2 = 0.73; P < 0.01) was observed
in the liver, where 18:1n-9 accumulation took place
when LO was included at 75 and 100% (Table 4). Liver
NL fatty acid composition reflected the effects of LO on
the elongation and desaturation processes (Bell et al.,
1993). The accumulation of 20:4n-3, the product of 6
desaturase, and the reduction of 20:4n-6 production are
probably indications of competitiveness of substrates
for the 5 desaturase (Bell et al., 1993). Thus, feeding
diets containing 75 or 100% LO notably increased the
accumulation of desaturated and elongated products
from 18:3n3 in the liver (Tables 4 and 5), in particular
the linear (R2 = 0.85; P < 0.001) and quadratic (R2 =
0.85; P < 0.001) effect of dietary LO inclusion level on
20:4n-3 accumulation in the liver.
Results presented in Tables 4 and 6 clearly indicated
that the proportion of 20:5n-3 and 22:6n-3 in salmon
liver and muscle triglycerides decreased accordingly
with the replacement of FO by LO. The proportion of
20:5n-3 decreased following a linear (R2 = 0.99; P <
0.001) and quadratic (R2 = 0.99; P < 0.001) pattern in
the muscle, whereas it followed a linear (R2 = 0.88; P <
0.001) decrease in the liver (Tables 4 and 6). Conversely,
2858
Menoyo et al.
g total fatty acids) of polar hepatic lipids
Item
No. of fish
Stearic acid (18:0)
Total SFA
Total MUFA
n-3:n-6 ratio
n-3 HUFAa
UIb
25
50
75
100
Pooled SE
P<F
5
2.12z
23.01z
6.22y
31.87z
1.72z
9.16x
0.10x
14.58x
1.17w
0.14z
3.49y
4.80x
0.21v
0.08y
0.12w
0.78x
12.00z
3.35z
31.78z
48.32z
10.86x
10.13z
48.03z
3.04z
5
1.54y
20.55xy
9.07z
31.51z
1.22y
10.33x
0.15xy
15.16xy
2.12x
0.13z
4.51z
6.77yz
1.75w
0.06y
0.86x
0.64x
9.19xy
2.84yz
30.41z
45.74y
12.75x
6.81y
43.93y
2.93xy
5
1.41y
21.55yz
6.59y
29.90yz
1.18y
9.83x
0.16xyz
14.18x
2.54x
0.10y
3.41y
6.05y
3.45x
0.08y
1.31y
0.94xy
10.03y
2.60y
30.53z
48.95z
12.21x
8.16y
45.42y
3.00yz
5
1.17x
19.71xy
6.72y
27.89y
1.14y
12.24y
0.27z
16.98yz
3.77y
0.10y
2.82y
6.68yz
5.75y
0.11y
2.03z
1.13y
8.11wx
2.33xy
27.79y
47.25yz
15.26y
7.11y
41.39x
2.86xy
5
0.67w
18.37x
6.11y
25.29x
0.74x
14.52z
0.22yz
18.34z
5.53z
0.05x
1.57x
7.15z
8.43z
0.26z
2.44z
2.47z
6.97w
1.83x
25.34x
47.75z
17.74z
6.72y
39.05w
2.79w
0.06
0.59
0.56
0.45
0.07
0.47
0.03
0.58
0.13
0.00
0.26
0.34
0.23
0.03
0.10
0.09
0.45
0.14
0.57
0.68
0.52
0.52
0.73
0.03
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
0.003
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
0.875
<0.001
<0.001
<0.001
<0.001
v,w,x,y,z
22:6n-3 decreased linearly (R2 = 0.98; P < 0.001) in the

muscle NL fraction (Table 6) and followed a linear (R2 =
0.74; P < 0.001) and a quadratic (R2 = 0.74; P < 0.01)
trend in the liver at the higher FO substitution levels
(Table 4). Regardless of the amount of 22:6n-3 provided
in the diet, this fatty acid was selectively deposited in
both tissues (Tables 4 and 6), and, according to Bell et
al. (2003b), this could be a consequence of the catabolic
complexity necessary to yield energy and a consequence
of the paramount role of 22:6n-3 as a cell membrane
component (Tables 5 and 7).
Increasing levels of LO in the diet promoted a different pattern of individual fatty acid accumulation in
liver and muscle PL and NL fractions (Table 8). Although total MUFA accumulated in the liver NL following a linear (R2 = 0.59; P < 0.001) and quadratic (R2 =
0.59; P < 0.03) fashion, there was a linear (R2 = 0.70; P
< 0.001) decrease in muscle NL. Conversely, increasing
substitution levels of dietary LO followed a different
pattern, with n-3 fatty acids increasing linearly (R2 =
0.88; P < 0.001) and quadratically (R2 = 0.88; P < 0.01)
in muscle NL as dietary LO increased; however, there
was a linear (R2 = 0.29; P < 0.005) decrease in n-3
concentrations in liver NL (Table 8). The proportion of
SFA tended to be diminished in the NL fraction of both
tissues, decreasing linearly in the muscle (R2 = 0.96; P
< 0.0001) and in a linear (R2 = 0.85; P < 0.001) and
quadratic (R2 = 0.85; P < 0.007) fashion in the liver
(Table 8). The fish tends to maximize the accumulation

of n-3 HUFA in the PL fraction of both tissues; however,
with the FO replacement, there was a progressive loss
of n-3 HUFA in both tissues that was more marked
in the liver than in the muscle at the high levels of
LO inclusion.
Previous studies performed with Atlantic salmon
have shown a lack of an effect of vegetable oils on fillet
quality measurements (Rosenlund et al., 2001b; Rra
et al., 2003) and organoleptic attributes (Obach et al.,
2001). One of the most important quality attributes in
fish is its nutritional value as a source of PUFA (Boggio
et al., 1985). Salmon rich in PUFA is, however, more
susceptible to lipid oxidation, yielding undesirable lipid
oxidation products that negatively affect quality characteristics. Although it is well known that unsaturated
fatty acids of the PL fraction are the main reason for
fish flesh peroxidation during storage (Monahan, 2000),
the susceptibility of flesh to oxidation also depends on
the balance of pro- and antioxidants (Lopez-Bote, 2000).
The effects of experimental diets on TBARS concentration in fish flesh stored under refrigeration for 9 d and
on the muscular contents of - and -tocopherols are
shown in Figures 2 and 3, respectively. A time effect
(P < 0.001) on TBARS formation was observed during
storage (Figure 2); however, no dietary effect was observed on TBARS concentration (P = 0.125). When feeding rainbow trout diets containing either herring oil or
2859
g of total fatty acids) of neutral intramuscular lipids
Item
No. of fish
Stearic acid (18:0)
Total SFA
Total MUFA
n-3:n-6 ratio
n-3 HUFAa
UIb
25
50
75
100
Pooled SE
P<F
5
5.43z
16.77z
3.80y
27.26z
7.43z
14.61w
2.59z
32.75y
4.40v
0.34z
0.82z
5.57v
1.43v
2.04z
1.70z
9.78z
3.48z
15.34z
33.99x
19.31x
6.11z
30.52z
2.25z
5
4.03y
15.11y
4.17z
24.17y
5.11y
19.32yz
2.43yz
34.31z
7.03w
0.28y
0.57y
7.89w
10.08w
1.33y
1.31y
5.97y
2.40y
11.06y
33.02x
24.04z
4.19y
21.61y
2.05y
5
3.40x
14.07x
4.09yz
22.27x
4.24x
17.68x
2.22y
30.50y
8.21x
0.23x
0.49x
8.93x
17.33x
1.25xy
1.28y
4.83x
1.92x
9.61x
37.66y
21.91xy
4.22y
19.08x
2.09y
5
2.71w
12.95w
4.23z
20.43w
3.27w
18.53xy
2.14y
29.96y
9.26y
0.20w
0.37w
9.83y
22.35y
1.08x
1.24y
3.47w
1.39w
7.45w
39.04y
22.85y
3.97y
15.61w
2.04y
5
1.48v
10.61v
4.20z
16.58v
1.87v
20.16z
1.74x
28.49x
11.05z
0.13v
0.21v
11.39z
29.67z
1.21xy
1.55z
1.83v
0.76v
4.74v
42.66z
24.01z
3.75x
11.78v
2.04y
0.08
0.21
0.08
0.32
0.13
0.60
0.07
0.54
0.17
0.03
0.01
0.15
0.51
0.06
0.04
0.22
0.05
0.29
0.56
0.64
0.15
0.54
0.02
<0.001
<0.001
0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
<0.001
v,w,x,y,z
b
Table 7. Effects of dietary fish and linseed oil combinations on selected fatty acids (g/
100 g of total fatty acids) of polar i.m. lipids
Item
No. of fish
Stearic acid (18:0)
Total SFA
Total MUFA
n-3:n-6 ratio
n-3 HUFAa
UIb
a
25
50
75
100
Pooled SE
P<F
5
2.29z
17.29z
3.24
23.39z
2.75z
9.33y
0.73
17.11
2.24w
0.12z
1.09z
3.55w
1.83w
0.45
1.28y
9.91z
3.27z
35.31z
52.31
11.62yz
15.47z
50.03z
3.25z
5
2.34z
17.74z
3.08
23.70z
2.53z
10.79yz
0.86
18.50
3.73wx
0.11yz
0.86yz
4.72wx
6.22wx
0.65
1.16y
8.31yz
2.89yz
32.78z
52.68
13.06yz
11.58yz
45.81z
3.23z
5
1.92z
16.73yz
3.70
22.81yz
2.06yz
9.06y
0.69
15.54
3.60x
0.10yz
1.01z
4.71xy
8.15x
0.58
1.16y
8.54yz
2.81yz
34.41z
56.55
10.98y
12.55yz
47.83z
3.25z
5
1.66yz
16.18yz
2.71
20.92yz
1.68yz
11.08yz
0.87
17.47
4.96y
0.10yz
0.64y
5.71y
13.82y
0.55
1.26y
6.97xy
2.15xy
28.94yz
55.36
13.51yz
9.77yx
41.00yz
3.05yz
5
0.97y
14.80y
3.76
19.74y
0.99y
12.63z
0.73
17.51
6.52z
0.07y
0.56y
7.16z
19.55z
0.67
1.86z
5.01x
1.55x
23.61y
54.83
14.99z
7.71x
34.61y
2.86y
0.20
0.50
0.50
0.73
0.24
0.73
0.06
0.66
0.26
0.01
0.08
0.25
1.06
0.05
0.05
0.56
0.20
1.56
1.40
0.88
0.88
2.12
0.07
<0.001
0.003
0.524
<0.001
0.001
0.028
0.493
0.292
<0.001
0.010
<0.001
<0.001
<0.001
0.176
<0.001
<0.001
<0.001
0.001
0.356
0.007
<0.001
<0.001
<0.001
Highly unsaturated fatty acids (HUFA) includes 20:3, 20:4, 20:5, 22:5, and 22:6.
Within a row, means lacking a common superscript letter differ, P < 0.05.
v,w,x,y,z
2860
Menoyo et al.
Table 8. Effects of different levels of dietary linseed oil (LO; % of total added fat) on fatty
acid classes (g/100 g of total fatty acids) in muscle and liver neutral lipids (NL) and polar
lipids (PL)a
P-value
LO
LO2
R2
Pooled SE
Linear
Quadratic
27.08
33.72
33.52
29.77
0.09
0.051
+0.028
0.277
+0.00063
+0.001
0.96
0.70
0.88
0.96
0.31
0.54
0.56
0.53
<0.001
<0.001
<0.001
<0.001
0.01
<0.001
Muscle PL
Saturated
n-3 HUFA
y = 24.09
y = 52
0.04
0.152
0.45
0.55
0.72
2.13
<0.001
<0.001
Liver NL
Saturated
Monounsaturated
n-3
n-3 HUFA
y
y
y
y
24.03
19.97
50.31
47.72
0.012
0.057
0.058
0.055
0.0009
+0.001
0.0018
0.85
0.59
0.29
0.79
0.75
1.85
1.51
2.05
<0.001
<0.001
0.005
<0.001
0.007
0.03
0.04
Liver PL
Saturated
Monounsaturated
n-3 HUFA
y = 31.94
y = 14.73
y = 47.66
0.011
0.023
0.081
0.0005
+0.0006
0.86
0.59
0.77
0.45
0.57
0.73
<0.001
<0.001
<0.001
0.009
0.02
Fatty acid
Intercept
Muscle NL
Saturated
Monounsaturated
n-3
n-3 HUFA
y
y
y
y
=
=
=
=
=
=
=
=
lard (approximately 10% DM), Boggio et al. (1985) failed

to detect differences in TBARS concentration in frozen
stored fillets, although higher TBARS were detected in
the fillet of fish fed lower amounts of -tocopheryl acetate (50 mg/kg of diet). Conversely, Olsen et al. (1999)
reported increasing TBARS values in response to increasing dietary PUFA levels in Arctic charr, which
indicated that supplying -tocopherol above requirements does not improve oxidative stability. Our laboratory previously reported higher TBARS values in the
flesh of Atlantic salmon fed a diet containing a rich n3 HUFA FO than when feeding diets formulated with
FO that was less rich in n-3 fatty acids (Menoyo et al.,
2002). When Atlantic salmon were marketed at approximately 5 kg, Menoyo et al. (2002) demonstrated that
Figure 2. Effect of dietary linseed oil inclusion level (0,

25, 50, 75, and 100% replacement of fish oil) on thiobarbituric acid reactive substances (TBARS; mol of malondialdehyde [MDA]/kg of wet tissue) concentration along
fish flesh stored under refrigeration for 9 d (P < 0.001 for
time effect).
inclusion of lower levels of -tocopheryl acetate in the

diet (170 mg/kg of feed, on average) was not sufficient
to palliate peroxidation effects between treatments.
Overall unsaturation and total n-3 HUFA concentrations were greater in i.m. lipids in fish receiving a diet
containing FO than in those receiving LO; therefore, a
higher oxidation would be expected.
In the present experiment, dietary oils provided not
only a different source of fatty acids but also a different
supply of natural antioxidants (Table 1), which might
affect susceptibility of tissues to lipid peroxidation (Jensen et al., 1998; Mortensen and Skibsted, 2000). Both
- and -tocopherol were greater (P < 0.02) in muscle
of fish fed the 100% LO diet, and muscle -tocopherol
Figure 3. Effect of increasing linseed oil (% replacement

of fish oil) in the feed of Atlantic salmon on muscle tissue
(mg/g of muscle) - (solid bars) and -tocopherol (open
bars) concentration. Values are on a wet-tissue basis. Bars
that do not have a common letter differ, P < 0.02.
concentration was almost five times greater in muscle

of fish fed 100% compared with 0% LO (Figure 3). Tocopherol has a greater antioxidant effect than other
vitamin E isoforms in Atlantic salmon tissue (Parazo
et al., 1998) and pork (Rey et al., 1998). Although tocopherol seems to be more active than -tocopherol
because of its tissue localization and mobilization (Parazo et al., 1998; Bell et al., 2000), additional research
is warranted to assess their potential as ante- and postmortem antioxidants in fish muscle.
Implications
Dietary linseed oil can totally replace fish oil in highenergy diets for Atlantic salmon weighing approximately 520 g without affecting fish performance and
fillet peroxidative stability. Nonetheless, fatty acid composition in muscle and liver was changed, leading to a
decrease in the concentration of n-3 highly unsaturated
fatty acids in fish flesh when diets with greater linseed
oil contents were fed.
Literature Cited
Alvarez, M. J., A. Diez, C. J. Lopez-Bote, M. Gallego, and J. M.
Bautista. 2000. Short-term modulation of lipogenesis by macronutrients in rainbow trout hepatocytes Br. J. Nutr. 84:110.
Bautista, J. M., A. Garrido-Pertierra, and G. Soler. 1988. Glucose-6phosphate dehydrogenase from Dicentrarchus labrax liver: Kinetic mechanism and kinetic of NADPH inhibition. Biochim.
Biophys. Acta 967:354363.
Bell, J. G., J. R. Dick, and J. R. Sargent. 1993. Effects of diets rich in
linoleic or -linolenic acid on phospholipid fatty acid composition
and eicosanoid production in Atlantic salmon (Salmo salar).
Lipids 28:819826.
Bell, J. G., R. J. Henderson, D. R. Tocher, F. McGhee, J. R. Dick, A.
Porter, R. P. Smullen, and J. R. Sargent. 2002. Substituting fish
oil with crude palm oil in the diet of Atlantic salmon (Salmo
salar) affects muscle fatty acid composition and hepatic fatty
acid metabolism. J. Nutr. 132:222230.
Bell, J. G., J. McEvoy, D. R. Tocher, F. McGhee, P. J. Campbell, and
J. R. Sargent. 2001. Replacement of fish oil with rapeseed oil
in diets of Atlantic salmon (Salmo salar) affects tissue lipid
composition and hepatocyte fatty acid metabolism. J. Nutr.
131:15351543.
Bell, J. G., J. McEvoy, D. R. Tocher, and J. R. Sargent. 2000. Depletion
of -tocopherol and astaxanthin in Atlantic salmon (Salmo salar)
affects autoxidative defense and fatty acid metabolism. J. Nutr.
130:18001808.
Bell, J. G., F. McGhee, P. J. Campbell, and J. R. Sargent. 2003a.
Rapeseed oil as an alternative to marine fish oil in diets of postsmolt Atlantic salmon (Salmo salar): Changes in flesh fatty acid
composition and effectiveness of subsequent fish oil wash out.
Aquaculture 218:515528.
Bell, J. G., A. H. McVicar, M. T. Park, and J. R. Sargent. 1991.
High dietary linoleic acid affects the fatty acid compositions of
individual phospholipids from tissues of Atlantic salmon (Salmo
salar): Association with stress susceptibility and cardiac lesion.
J. Nutr. 121:11631172.
Bell, J. G., D. R. Tocher, R. J. Henderson, J. R. Dick, and V. O.
Crampton. 2003b. Altered fatty acid compositions in Atlantic
salmon (Salmo salar) fed diets containing linseed and rapeseed
oils can be partially restored by a subsequent fish oil finishing
diet. J. Nutr. 133:27932801.
Boggio, S. M., R. W. Hardy, J. K. Babbitt, and E. L. Brannon. 1985.
The influence of dietary lipid source and alpha-tocopheryl ace-
2861
tate level on product quality of rainbow trout (Salmo gairdneri).

Bradford, M. M. 1976. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of
protein-dye binding. Anal. Biochem. 72:248254.
Caballero, M. J., A. Obach, G. Rosenlund, D. Montero, M. Gisvold,
and M. S. Izquierdo. 2002. Impact of different dietary sources
on growth, lipid digestibility, tissue fatty acid composition and
histology of rainbow trout, Oncorhynchus mykiss. Aquaculture
214:253271.
FAO. 2002. The State of World Fisheries and Aquaculture. Rome
Italy. Publication Division, Food and Agricultural Organization
of the United Nations (FAO), Viale delle Terme di Caracalla,
00100 Rome, Italy.
Fryland, L., L. Madsen, K. M. Eckhoff, . Lie, and R. K. Berge.
1998. Carnitine palmitoyltransferase I, carnitine palmitoyltransferase II, and acyl-CoA oxidase activities in Atlantic salmon
(Salmo salar). Lipids 33:923930.
Grisdale-Helland, B., B. Ruyter, G. Rosenlund, A. Obach, S. J. Helland, M. Gisvold, H. Standal, and C. Rsj. 2002. Influence of
high contents of dietary soybean oil on growth, feed utilization,
tissue fatty acid composition, heart histology and standard oxygen consumption of Atlantic salmon raised at two temperatures.
Jensen, C., E. Birk, A. Jokumsen, L. H. Skibsted, and G. Bertelsen.
1998. Effect of dietary levels of fat, -tocopherol and astaxanthin
on colour and lipid oxidation during storage of frozen rainbow
trout (Oncorhynchus mykiss) and during chill storage of smoked
trout. Z. Lebensm Unters Forsch A 207:189196.
Kiessling, K. H., and A. Keissling. 1993. Selective utilization of fatty
acids in rainbow trout (Oncorhynchus mykiss Walbaum) red
muscle mitochondria. Can. J. Zool. 71:248251.
Lopez-Bote, C. J. 2000. Dietary treatment and quality characteristics
in mediterranean meat products. Pages 345366 in Antioxidants
in Muscle Foods. E. A. Decker, C. Faustman, and C. J. LopezBote, eds. Wiley & Sons, Inc., New York, NY.
Lopez-Bote, C. J., A. I. Rey, M. Sanz, J. I. Gray, and D. J. Buckley.
1997. Dietary vegetable oils and -tocopherol reduce lipid oxidation in rabbit muscle. J. Nutr. 127:11761182.
Marmer, W. N., and R. J. Maxwell. 1981. Dry column method for the
quantitative extraction and simultaneous class separation of
lipids from muscle tissue. Lipids 16:365371.
Mckenzie, D. J., D. A. Higgs, B. Dosanjh, G. Deacon, and D. J. Randall.
1998. Dietary lipid composition influences swimming performance in Atlantic salmon (Salmo salar) in seawater. Fish Physiol. Biochem. 19:111122.
Menoyo, D., M. S. Izquierdo, L. Robaina, R. Gines, C. J. Lopez-Bote,
and J. M. Bautista. 2004. Adaptation of lipid metabolism, tissue
composition and flesh quality in gilthead sea bream (Sparus
aurata) to the replacement of dietary fish oil by linseed and
soyabean oils. Br. J. Nutr. 92:4152.
Menoyo, D., C. J. Lopez-Bote, J. M. Bautista, and A. Obach. 2002.
Herring vs. anchovy fish oils in salmon feeding. Aquat. Living
Resour. 15:217223.
Menoyo, D., C. J. Lopez-Bote, J. M. Bautista, and A. Obach. 2003.
Growth, digestibility and fatty acid utilization in large Atlantic
salmon (Salmo salar) fed varying levels of n-3 and saturated
fatty acids. Aquaculture 225:295307.
Monahan, F. J. 2000. Oxidation of lipids in muscle foods: Fundamentals and applied concerns. Pages 324 in Antioxidants in Muscle
Foods. E. A. Decker, C. Faustman, and C. J. Lopez-Bote. eds.
Wiley & Sons, Inc., New York, NY.
Morris, T. R. 1999. Experimental Design and Analysis in Animal
Sciences. CAB Int., Oxford, UK.
Mortensen, A., and L. H. Skibsted. 2000. Antioxidant activity of carotenoids in muscle foods. Pages 6185 in Antioxidants in Muscle
Foods. E. A. Decker, C. Faustman, and C. J. Lopez-Bote, eds.
Wiley & Sons, Inc., New York, NY.
NRC. 1993. Pages 1315 in Nutrient Requirements of Fish. Natl.
Acad. Press, Washington, DC.
2862
Menoyo et al.
. Bendiksen, G. Rosenlund, and M. Gisvold. 2001.

Obach, A., E. A
Impact of dietary lipid source on muscle fatty acid composition
and sensory evaluation of Atlantic salmon (Salmo salar L.).
Pages 391392 in Farmed Fish Quality. S. C. Kestin and P. D.
Warriss, eds. Fishing News Books, Blackwell Sci. Ltd., Oxford, UK.
Olsen, R. E., E. Lvaas, and . Lie. 1999. The influence of temperature, dietary polyunsaturated fatty acids, -tocopherol and
spermine on fatty acid composition and indices of oxidative stress
in juvenile Arctic char (Salvelinus alpinus L.). Fish Physiol.
Biochem. 20:1329.
Parazo, M. P. M., S. P. Lall, J. D. Castell, and R. G. Ackman. 1998.
Distribution of - and -tocopherols in Atlantic salmon (Salmo
salar) tissues. Lipids 33:697704.
Regost, C., J. Arzel, J. Robin, G. Rosenlund, and S. J. Kaushik. 2003.
Total replacement of fish oil by soybean or linseed oil with a
return to fish oil in turbot (Psetta maxima) 1. Growth performance, flesh fatty acid profile, and lipid metabolism. Aquaculture 217:465482.
Rey, A. I., B. Isabel, R. Cava, and C. J. Lopez-Bote. 1998. Dietary
acorns provide a source of gamma-tocopherol to pigs raised extensively. Can. J. Anim. Sci. 78:441443.
Rey, A. I., J. P. Kerry, P. B. Lynch, C. J. Lopez-Bote, D. J. Buckley,
and P. A. Morrissey. 2001. Effect of dietary oils and -tocopheryl
acetate supplementation on lipid (TBARS) and cholesterol oxidation in cooked pork. J. Anim. Sci. 79:12011208.
. Bendiksen, M. Gisvold, and B. Ruyter.
Rosenlund, G., A. Obach, E. A
2001a. Effect of dietary fatty acid composition on Atlantic salmon
(Salmo salar) when replacing fish oils with vegetable oils. Pages
403404 in Farmed Fish Quality. S. C. Kestin and P. D. Warriss,
eds. Fishing News Books, Blackwell Sci. Ltd., Oxford, UK.
Rosenlund, G., A. Obach, M. G. Sandberg, H. Standal, and K. Tveit.
2001b. Effect of alternative lipid sources on long-term growth
performance and quality of Atlantic salmon (Salmo salar). Aquacult. Res. 32(Suppl. 1):17.
Rra, A. M. B., C. Regost, and J. Lampe. 2003. Liquid holding capacity,
texture and fatty acid profile of smoked fillets of Atlantic salmon
fed diets containing fish oil or soybean oil. Food Res. Int.
36:231239.
Sanz, M., C. Lopez-Bote, D. Menoyo, and J. M. Bautista. 2000. Abdominal fat deposition and fatty acid synthesis are lower and oxidation is higher in broiler chickens fed diets containing unsaturated rather than saturated fat. J. Nutr. 130:30343037.
Sukhija, P. S., and D. L. Palmquist. 1988. Rapid method for determination of total fatty acid content and composition of feedstuffs
and feces. J. Agric. Food Chem. 36:12021206.
Tocher, D. R., J. G. Bell, J. R. Dick, R. J. Henderson, F. McGee,
D. Michell, and P. C. Morris. 2000. Polyunsaturated fatty acid
metabolism in Atlantic salmon (Salmo salar) undergoing parrsmolt transformation and the effects of dietary linseed and rapeseed oils. Fish Physiol. Biochem. 23:5973.
Tocher, D. R., J. G. Bell, P. MacGlaughlin, F. McGee, and J. R. Dick.
2001. Hepatocyte fatty acid desaturation and polyunsaturated
fatty acid composition of liver in salmonids: Effects of dietary
vegetable oil. Comp. Biochem. Biophys. B130:257270.
Thompson, K. D., M. F. Tatner, and R. J. Anderson. 1996. Effects of
dietary (n-3) and (n-6) polyunsaturated fatty acid ratio on the
immune response of Atlantic salmon, Salmo salar L. J. Aqua.
Nutr. 2:2131.
Torstensen, B. E., . Lie, and L. Fryland. 2000. Lipid metabolism
and tissue composition in Atlantic salmon (Salmo salar L.).
Effects of capelin oil, palm oil, and oleic acid-enriched sunflower
oil as dietary lipid sources. Lipids 35:653664.
Zammit, V. A. 1999. The malonyl-CoA-long chain acyl-CoA axis in
the maintenance of mammalian cell function. Biochem. J.
343:505515.

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Effect of dietary fish oil substitution with linseed oil on the performance, tissue

fatty acid profile, metabolism, and oxidative stability of Atlantic salmon1,2

ABSTRACT: The objective of this experiment was to

the polar lipid fraction (PL) were unaffected (P = 0.356)

J. Anim. Sci. 2005. 83:28532862

mental effects on growth. Nonetheless, use of vegetable

Table 1. Composition of experimental diets

LT = low temperature (71.8 CP/8.4 moisture)

binations for Atlantic salmon on fatty acid composition

Materials and Methods

Biological Parameters and Sample Collection

recorded. Then, fish were eviscerated, and the weight

Chemical Analyses of Flesh Samples and Lipid

Linseed oil in salmon feeding

Table 2. Fatty acid composition (%) of experimental diets

Mitochondrial Preparations, Soluble Extracts,

A repeated measure mean test was used to compare

Results and Discussion

adiposity (Bell et al., 2001, 2002), whereas LO had no

concentration of total SFA, and a higher concentration

Figure 1. Effect of increasing linseed oil (% replacement

Linseed oil in salmon feeding

such as changes in 9 desaturase activity, also might

22:6n-3 decreased linearly (R2 = 0.98; P < 0.001) in the

(Table 8). The fish tends to maximize the accumulation

Linseed oil in salmon feeding

lard (approximately 10% DM), Boggio et al. (1985) failed

Figure 2. Effect of dietary linseed oil inclusion level (0,

inclusion of lower levels of -tocopheryl acetate in the

Figure 3. Effect of increasing linseed oil (% replacement

Linseed oil in salmon feeding

concentration was almost five times greater in muscle

tate level on product quality of rainbow trout (Salmo gairdneri).

. Bendiksen, G. Rosenlund, and M. Gisvold. 2001.

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