Fatty acid content and potential health benefits of consuming gilthead sea bream ( Sparus aurata ) and sea bass ( Dicentrarchus labrax )

Abstract

The fatty acid composition of wild sea bass (Dicentrarchus labrax L.) and gilthead sea bream (Sparus auratus L.) were compared with gas chromatography. These two species are widely cultivated in Europe and represent a significant portion of consumed fish in the region. The aim of the present work was to compare the nutritional value of fatty acids in the flesh of wild sea bass and sea bream. Significant differences were observed in the saturated and poly-unsaturated fatty acid content. The presence of lauric, myristic and palmitic acids in the flesh of sea bream in quantities far exceeding those in sea bass make sea bream less suitable for preventing cardiovascular diseases. The poly-unsaturated n-3 fatty acids with both anti-atherogenetic and anti-inflammatory action in sea bass surpass those of sea bream by a total of 30%. Sea bass also contains 60% more C22:6n-3. Compared to sea bream, sea bass appears to be more suitable for the diet of people suffering from cardiac diseases, angiopathy, inflammations and Alzheimer's disease.

Full text

0139–3006/$ 20.00 © 2011 Akadémiai Kiadó, Budapest
Acta Alimentaria, Vol. 40 (1), pp. 45–51 (2011)
DOI: 10.1556/AAlim.40.2011.1.7
FATTY ACID CONTENT AND POTENTIAL HEALTH BENEFITS
OF CONSUMING GILTHEAD SEA BREAM (SPARUS AURATA) AND
SEA BASS (DICENTRARCHUS LABRAX)
D. LENASa*, E. PAPADIMITRIOUb, C. BITCHAVAc and C. NATHANAILIDESa
aDepartment of Ichthyology and Fisheries,
Technological Educational Institute of Epirus, 46100 Igoumenitsa. Greece
bDepartment of Nursing, School of Health Social Welfare,
Technological Educational Institute of Epirus, 45500 Ioannina. Greece
c Laboratory Of Ichthyology and Aquatic Animal Health, Faculty of Veterinary Medicine,
School of Health Sciences, University of Thessaly, Greece 43100, Karditsa. Greece
(Received: 15 July 2009; accepted: 2 July 2010)
The fatty acid composition of wild sea bass (Dicentrarchus labrax, L.) and gilthead sea bream (Sparus auratus, L.)
were compared with gas chromatography. These two species are widely cultivated in Europe and represent a
signifi cant portion of consumed fi sh in the region. The aim of the present work was to compare the nutritional value
of fatty acids in the fl esh of wild sea bass and sea bream. Signifi cant differences were observed in the saturated and
poly-unsaturated fatty acid content. The presence of lauric, myristic and palmitic acids in the fl esh of sea bream in
quantities far exceeding those in sea bass make sea bream less suitable for preventing cardiovascular diseases. The
poly-unsaturated n-3 fatty acids with both anti-atherogenetic and anti-infl ammatory action in sea bass surpass those
of sea bream by a total of 30%. Sea bass also contains 60% more C22:6n-3. Compared to sea bream, sea bass appears
to be more suitable for the diet of people suffering from cardiac diseases, angiopathy, infl ammations and Alzheimer’s
disease.
Keywords: seafood, fi sh, nutritional value, Dicentrarchus labrax, cardiovascular
The nutritional value of fi sh is attributed not only to amino-acids, trace elements, and vitamins
it contains, but also, and most importantly, to the quantity and quality of fatty acids (SARGENT
1997; BUCHTOVA et al., 2008; 2009). For example, increased consumption of food with high
ratio of n-6/n-3 fatty acids leads to thrombosis and infl ammatory reactions in the tissue
(SARGENT et al., 1999), while high contents of n-3/n-6 have, among other benefi cial effects for
the prevention of cardiovascular diseases, antithrombotic and anti-infl ammatory effects
(KELI et al., 1994; KRIS-ETHERTON et al., 2003; DIN et al., 2004). The balance between n-3
and n-6 is also important for homeostasis and the normal development of the body (KRIS-
ETHERTON et al., 2000; SARGENT, 1997), while also infl uences receptor’s function and the
activity of many membrane-bound enzymes (JUMPSEN & CLANDININ, 1995). The best n-3/n-6
ratio has not been exactly determined for superior mammals. The ratio of 1:5 (n-3/n-6) is
today considered to be near ideal universally, though for fi sh this relation can be smaller
(SARGENT et al., 1999). More specifi cally, arachidonic acid (AA) which is an omega-6 essential
fatty acid, eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) which both are
omega-3 fatty acids, are considered essential for the structure and operation of cell membranes
(SARGENT et al., 1993; 1995; SARGENT, 1997). The thrombotic PAF (Platelet Activating Factor)
* To whom correspondence should be addressed.
Phone: 2665049882; fax: 2665049886; e-mail: lenasds@teiep.gr

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LENAS et al.: CARDIOVASCULAR HEALTH ASPECTS OF WILD SEA BASS
role of lipids have been ascribed mainly to polar lipids, while the antithrombotic anti-PAF
are ascribed to neutral lipids and more specifi cally to minor lipid compounds (vitamin
E, carotenoids, phytosterols, etc.) (NOMIKOS et al., 2006; NASOPOULOU et al., 2007) which are
found predominantly in wild marine fi sh (including sea bass, sea bream, etc.).
Omega-3 dietary intake has several health benefi ts (MANCARDI et al., 2009) and can
limit arteriosclerosis in vivo when combined with PAF competitors (MAYER et al., 2002).
The content in lipids and minor lipid compounds in the fl esh of wild fi sh depends on their
feeding habits in the natural environment they live. This content also characterises their
nutritional value. Seafood and mainly fi sh are the most important natural foods that supply
the human organism with n-3 fatty acids and minor lipid compounds. Healthy food
consumption patterns should be encouraged in the general population with special attention
to particular risk groups including pregnant women and the elderly (ELMADFA & MEYER,
2009).
The aim of this study was to rate the nutritional value of fatty acids in the edible fl esh of
two wild fi sh that are consumed widely in Europe (PIENIAK et al., 2009), the sea bass
(Dicentrarchus labrax) and the gilthead sea bream (Sparus aurata). This information is
crucial for nutritionists and for the food industry. These two fi sh species are consumed fresh
and on several occasions are sold fi lleted. The fi sh by-products generated by the fi lleting can
be used by the food and the pharmaceutical industry for extracting EPA and DHA (OLIVEIRA
et al., 2009).
1. Materials and methods
1.1. Biological material
We examined wild European sea bass and gilthead sea bream specimens, all randomly
selected, originating from the delta of River Kalamas in the coastal area of the Ionian Sea
(Prefecture of Thesprotia, western Greece). The ages of the fi sh were 2+ to 3+.
Initially, one hundred wild fi sh (forty four sea bass and fi fty six sea bream) were collected
alive in May 2008 using fi shing nets. The captured fi sh were anaesthetized in a solution of
150 mg l–1 clove oil (PERDIKARIS et al., 2010) and those of commercial size were killed in an
ice cold bath and immediately transported to the lab in ice. Subsequently, the fi sh were
cleaned with fresh water. Body weight and total length were measured to the closest mg and
mm, respectively, before they were scaled, skinned, de-headed, eviscerated and fi lleted. All
body parts of each fi sh were weighted, marked, vacuum packed in plastic bags and stored in
–30 oC before the next stage of the study.
1.2. Age determination of the specimens
The ages of the fi sh were determined in the Department of Ichthyology and Fisheries of
the Technological Educational Institute of Epirus, based on the annual rings of the alcohol-
infused otoliths (PANELLA, 1971). For this reason, an inverted stereoscope with 50 ×
magnifi cation was used and the age was determined twice independently. Differential
determination of age led to the exclusion of the respective fi sh from the sample. In the same
manner, fi sh with less clear rings were also excluded. As a result from the hundred fi sh
examined, only thirty sea bass end forty sea bream were found to be between 2+ and 3+ age
groups.

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LENAS et al.: CARDIOVASCULAR HEALTH ASPECTS OF WILD SEA BASS
1.3. Determination of total lipids and fatty acids
Flesh samples were defrosted and chopped. Subsequently the edible parts of each fi sh were
homogenized in a mechanical homogenizer with metallic blades at low temperature (ice bath)
for one minute. Finally, twenty grams were used from the homogenate. The homogenized
brains (in pairs from each group of fi sh) were defrosted before the analysis. Lipid extraction
was performed by the BLIGH and DYER (1959) method as modifi ed by KINSELLA and co-
workers (1977), using chloroform and methanol in a 2:1 ratio. Subsequently, fatty acids were
methyl-esterifi ed with a 12% boron trifl uoride methanol solution (BF3-MeOH) (FOLCH et al.,
1957). Methyl-esters were obtained with normal hexane (METCALFE et al., 1966). The analysis
was performed using gas chromatography (Model GC-17A; Shimadzu, Kyoto, Japan) with a
capillary column and an ionised fl ame detector (TRACETM TR-FAME GC Column, Thermo
Fisher Scientifi c Inc.) as well as an automatic sampler (HT 310A, HTA). Pure helium of 82
KPa fl ow, air of 50 KPa fl ow and hydrogen of 60 KPa fl ow were used for the analysis under
the following conditions: the initial temperature was 150 oC for 5 min, followed by a 5 oC
min–1 increase to 170 oC for 10 min and then another 5 oC min–1 increase to until 220 oC for 20
min. The identifi cation of fatty acids (methyl-esters) was performed by comparing the
impressed peaks in special PC programme with Qalmix Fish (89-5550) and Methyl
Dodecanoate (20-1200) standard fatty acids (Larodan Fine Chemicals AB).
1.4. Statistical analysis
Statistical analysis (mean values, standard deviation and proportions) was performed using
Excel 2003 (Microsoft). T-tests were applied after variability comparison by F tests.
2. Results and discussion
The total lipids in the 100 grams of fi sh fl esh were found to be 1.55±0.35 in sea bass and
3.80±0.31 in sea breams.
With the aid of gas chromatography we identifi ed 30–45 fatty acids in the fl esh of both
categories of fi sh, from which only 18 were identifi ed based on standard fatty acids
(Table 1).
The saturated fatty acids were found to be 26.06% in sea bass and 27.46% in sea bream
with the predominant fatty acid in both cases being C16:0, followed by C18:0 and C14:0.
Mono-unsaturated fatty acids were identifi ed (at 37.83% and 37.42% respectively) with
C18:1 being prevalent in both cases, followed by C22:1 and C16:1.
The poly-unsaturated fatty acids (Fig. 1) found were 28.57% in sea bass compared to
22.82% in sea bream, with the dominant fatty acid in both cases being C22:6n-3 and followed
in sea bass by C18:2n-6 and C20:4n-6, while in sea bream by C22:50n-3, C20:4n-6 and
C18:2n-6.
The n-3 and n-6, as well as the ratio n-3/n-6 were higher in sea bass. C20:5n-3 (EPA)
was not detected in sea bass, while in sea bream it was detected in small quantities, which
indicates that in this early stage of the fi shes’ development the EPA is probably converted
very rapidly into C22:6n-3 for the needs of the cell membranes.
The main difference observed in the qualitative and quantitative constitution of the
edible fl esh of the fi sh examined was in saturated and poly-unsaturated fatty acids. So the
presence of lauric (C12:0), myristic (C14:0) and palmitic (C16:0) acids in the fl esh of sea

48
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LENAS et al.: CARDIOVASCULAR HEALTH ASPECTS OF WILD SEA BASS
bream and their higher quantities compared to those of sea bass makes the fl esh of gilthead
sea bream more harmful, both in terms of quality and quantity, for the health of consumers
and especially those suffering from cardiovascular diseases. These saturated fatty acids are
associated with hypercholesterolaemia and atherogenesis (KATHLEEN & ESCOTT, 2000).
The muscle content of gilthead sea bream is about twofold higher that sea bass.
Consequently, consumption of a serving of 100 g from the gilthead sea bream provides the
consumer double quantities of the less “desirable” dietary fatty acids C12:0, C14:0, and
C16:0, compared to sea bass. Furthermore, compared to gilthead sea bream, sea bass exhibits
increased content of PUFA n-3 fatty acids with well known anti-atherogenetic and anti-
Table 1. Fatty acid profi les of total lipids in the fl esh of wild sea bass (Dicentrarchus labrax) and gilthead sea
bream (Sparus aurata) (% of total fatty acids)a
Fatty acid Sea bass
N=10
Sea bream
N=10
P
t-test
C12:0 0.00±0.00 0.19±0.02 ***
C14:0 2.34±0.11 2.28±0.04 NS
C15:0 0.64±0.03 0.39±0.03 ***
C16:0 17.69±0.40 18.89±0.17 ***
C18:0 5.40±0.62 5.70±0.07 NS
Total saturated 26.06±1.14 27.46±0.32 **
C16:1 n-7 (9C) 5.72±0.28 6.23±0.07 ***
C18:1 n-9 (9C) 20.01±1.94 17.55±0.27 **
C18:1 n-7 (11C) 4.55±0.47 3.98±0.07 **
C20:1 n-9 (11C) 0.00±0.00 1.06±0.04 ***
C22:1 n-9 (13C) 7.55±0.36 8.60±0.29 ***
Total monounsaturated 37.83±2.97 37.42±0.59 NS
C18:2 n-6 4.42±0.29 2.63±0.04 ***
C18:3 n-3 1.53±0.03 1.18±0.10 ***
C18:4 n-3 0.62±0.02 1.01±0.07 ***
C20:4 n-6 3.63±0.23 3.42±0.07 *
C20:5 n-3 0.00±0.00 0.27±0.04 ***
C22:4 n-6 0.00±0.00 0.94±0.09 ***
C22:5 n-3 2.96±0.13 3.87±0.05 ***
C22:6 n-3 15.41±0.63 9.52±0.06 ***
Total polyunsaturated 28.57±1.05 22.82±0.51 ***
Others 7.5 12.30
Total n-3 fatty acids 20.52±0.54 15.83±0.31 ***
Total n-6 fatty acids 8.05±0.51 6.99±0.20 ***
Ratio n-3/n-6 2.55±0.10 2.27±0.03 ***
Ratio EPA/DHA 0.0±0.0 0.03±0.00 ***
aMean ±SD; N: Number of samplings; NS: Non signifi cant
*P< 0.05, **P<0.01, ***P<0.001

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LENAS et al.: CARDIOVASCULAR HEALTH ASPECTS OF WILD SEA BASS
infl ammatory effects such as decreasing concentration and oxidation of lipids and platelets,
deterrence of arrhythmias (BRAWN & HU, 2001), the suspension of the effects of synthetases
for the production of thromboxane (A2) (ABEYWARDENA et al., 1991) are in higher concentration
in sea bass compared to sea bream by a total of 30%, with the desirable levels for those with
a heart condition (score) >7.2 g/daily (SIMON et al., 1995) and average intake 0.45 g/daily
(VAN GELDER et al., 2007). The C22:6n-3 fatty acid was more abundant in sea bass (60%
compared to gilthead sea bream). This fatty acid is known to decrease VCAM-l in the vasal
endothelium and the adherence of leukocytes (HOLVOET, 1999; HU et al., 2001), in addition,
this fatty acid competes with linoleic acid in the arachidonic acid pathway, and reduces the
metabolism of AA to prostaglandin E2 and thromboxane A2, which are implicated in the
process of infl ammation. Furthermore, C22:6n-3 fatty acid decreases LR11 in the brain, a
protein that promotes Alzheimer’s disease (AD), (MA et al. 2007), and prevents the production
and concentration of b-amiloide (LIM et al., 2005). The n-3/n-6 ratio was also higher and thus
more benefi cial (RUSSO, 2009) in sea bass fi lets. The combination of sea bass with other plant
foods rich in C18: 2 fatty acids (vegetables, cereals) in the human diet offers a better and
healthier n-3/n-6 ratio, compared to the n3/n6 ratio of gilthead sea bream. Wild sea bass thus
appears more suitable than sea bream for the nutritional requirements of people suffering
from cardiopathy, angiopathy, infl ammations and Alzheimer’s disease.
3. Conclusions
The results of the present work indicate that compared to gilthead sea bream, the fatty acid
composition of sea bass exhibits signifi cantly higher concentration of fatty acids with well
known health benefi ts. On the contrary the fatty acid composition of sea bream is characterised
by a high content of saturated fatty acids such as lauric (C12:0), myristic (C14:0) and palmitic
(C16:0) acid, which are associated with hypercholesterolaemia and atherogenesis. In the
Fig. 1. Percentage composition of polyunsaturated fatty acids composition of wild fi sh sea bass and sea bream.
: sea bass; : sea bream

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same manner, the n-3/n-6 ratio of sea bass and gilthead sea bream, indicates that compared to
sea bream, sea bass is more appropriate for the diet of people suffering from cardiac diseases,
angiopathies, infl ammations and Alzheimer’s disease.
References
ABEYWARDENA, M.Y., MCLENNAN, P.L. & CHARNOCK, J.S. (1991): Differential effects of dietary fi sh oil on myocardial
prostaglandin I2 and Thromboxane A2 production. Am. J. Physiol., 260, H379-H385.
BLIGH, E.G. & DYER W.J. (1959): A rapid method of total lipid extraction and purifi cation. Can. J. Biochem. Physiol.,
37, 911–917.
BRAWN, A.A. & HU, F.B. (2001): Dietary modulation of endothelial function: implications for cardiovascular disease.
Am. J. clin. Nutr., 73, 673–686.
BUCHTOVÁ, H., SVOBODOVÁ, Z., KOCOUR, M. & VELÍŠEK, J. (2008): Chemical composition of edible parts of three-year-
old experimental scaly crossbreds of common carp (Cyprinus carpio, Linnaeus 1758). Acta Alimentaria, 37,
311–322.
BUCHTOVÁ, H., SVOBODOVÁ, Z., KOCOUR, M. & VELÍSEK, J. (2009): Amino acid composition in fi llets of mirror
crossbreds common carp (Cyprinus carpio). Acta Vet. Brno, 78, 337–344.
DIN, J.N., NEWBY, D.E. & FLAPAN, A.D. (2004): Omega 3 fatty acids cardiovascular disease- fi shing for a natural
treatment. Br. Med. J., 328, 30–35.
ELMADFA, I. & MEYER, A. (2009): Trends in nutrition in Europe. Acta Alimentaria, 38, 153–159.
FOLCH J., LESS, M. & STANLEY, G.H.S. (1957): A simple method for the isolation and purifi cation of total lipids from
animal tissues. J. biol. Chem., 226, 457–509.
HOLVOET, P. (1999): Endothelial Dysfunction, oxidation of low-density lipoprotein, and cardiovascular disease.
Therapeutic apheresis, 3, 287–293.
HU, F.B., MANSON, J.E. & WILLET, W.C. (2001): Types of dietary fat and risk of coronary heart disease: A critical
Review. J. Am. Coll. Nutr., 1, 5–19.
JUMPSEN, J. & CLANDININ, M.T. (1995): Brain development: Relationship to dietary lipid and lipid metabolism.
The American Oil Chemists Society, Illinois, p. 119.
KATHLEEN, M.L. & ESCOTT, S.S. (2000): Krause’s food, nutrition and diet therapy. 10th ed. W.B. Saunders, Philadelphia,
p. 327.
KELI, S.O., FESKENS, E.J.M. & KROMHOUT, D. (1994): Fish consumption and risk of stroke: The Zutphen study.
Stroke, 25, 328–332.
KINSELLA, J.E., SHIMP, J.L. & WEIHRAUCH, J. (1977): Fatty acid content and composition of freshwater fi nfi sh. J. Am.
Oil Chem. Soc., 54, 424–429.
KRIS-ETHERTON, P.M., TAYLOR, D.S., YO-POTH, S., HUTH, P., MORLARTY, K., FISHELL, V., HANGROVE, R.L., ZHAO, G. &
ETHERTON, T.D. (2000): Polyunsaturated fatty acids in the food chain in the United States. Am. J. clin. Nutr.,
71, 179S–88S.
KRIS-ETHERTON, P.M., HARRIS, W.S. & APPEL, L.J. (2003): Fish consumption fi sh oil, ω-3 fatty acids, and cardiovascular
disease. Arterioscl. Thromb. vascular Biol., 23, 20–31.
LIM, G.P., CALON, F., MORIHARA, Y., YANG, F., TETER, B., UBEDA, O., SALEM, N., FRAUTSCHY, S.A. & COLE, G.M.
(2005): A diet enriched with omega-3 fatty acid docosahexaenoic acid reduces amyloid burden in an aged
Alzheimer mouse model. J. Neurosci., 23, 3032–3040.
MA, Q.L., TETER, B., UBEDA, O.J., MORIHARA, T., DHOOT, D., NYBY, M.D., TUCK, M.L., FRAUTSCHY, S.A. & COLE,
G.M. (2007): Omega-3 fatty acid docosahexaenoic acid increases SorLa/LR11, a sorting protein with reduced
expression in sporadic Alzheimer’s disease (AD): Relevance to AD prevention. J. Neurosci., 27, 14299–
14307.
MANCARDI, D., TULLIO, F., CRISAFULLI, A., RASTALDO, R., FOLINO, A., PENNA, C. & PAGLIARO, P. (2009): Omega 3 has
a benefi cial effect on ischemia/reperfusion injury, but cannot reverse the effect of stressful forced exercise.
Nutr. Metab. Cardiovasc. Dis., 19, 20–26.
MAYER, K., MARFELS, M., MUHLY-REINHOLZ, M., GOKORSCH, S., ROSSEAU, S., LOHMEYER, J. SCHWARZER, N., KRÜLL, M.,
SUTTORP, N., GRIMMINGER, F. & SEEGER, W. (2002): ω-3 Fatty acids suppress monocyte adhesion to human
endothelial cells: role of endothelial PAF generation. Am. J. Physiol. Heart Circ. Physiol., 283, H811-H818.
METCHALFE, L.D., SCHMITZ, A.A. & PELKA, J.R. (1966): Rapid preparation of fatty acids esters from lipids for gas
chromatography analysis. Ann. Chem., 38, 524–535.

51
Acta Alimentaria 40, 2011
LENAS et al.: CARDIOVASCULAR HEALTH ASPECTS OF WILD SEA BASS
NOMIKOS, T., KARANTONIS, H.C., SKARVELIS C., DEMOPOULOS, C.A. & ZABETAKIS, I. (2006): Platelet activating properties
of lipid fractions obtained from raw and fried fi sh samples. Fd Chem., 96, 29–35.
NASOPOULOU, C., NOMIKOS, T., DEMOPOULOS, C.A. & ZABETAKIS, I. (2007): Comparison of antiatherogenic properties
of lipids obtained from wild and cultured sea bass (Dicentrarchus labrax) and gilthead sea bream (Sparus
aurata). Fd Chem., 100, 560–567.
OLIVEIRA, G.F., DESCHAMPS, F.C. & PESSATTI, M.L. (2009): The fatty acid composition of Brazilian fi shery by-
products. It. J. Fd Sc., 21, 231–239.
PANELLA, G. (1971): Fish otoliths: Daily growth layers and periodical patterns. Science, 173, 1124–1126.
PERDIKARIS, C., NATHANAILIDES, C., GOUVA, E., GABRIEL, U.U., BITCHAVA, C., ATHANASSOPOULOU, F., PASCHOU, A. &
PASCHOS I. (2010): Size-related effectiveness of Clove oil as anaesthetic for rainbow trout (Oncorhynchus
mykiss Walbaum, 1792) and goldfi sh (Carassius auratus Linnaeus, 1758). Acta Vet. Brno, 79, (in press)
PIENIAK, Z., VERBEKE, W., BRUNSØ, K., SCHOLDERER, J. & OLSEN, S. (2009): Comparison between Polish and Western
European fi sh consumers in their attitudinal and behavioural patterns. Acta Alimentaria, 38, 179–192.
RUSSO, G.L. (2009): Dietary n−6 and n−3 polyunsaturated fatty acids: From biochemistry to clinical implications in
cardiovascular prevention. Biochem. Pharmacol., 77, 937–946.
SARGENT, J.R. (1997): Fish oils and human diet. Br. J. Nutr., 78, 5–13.
SARGENT, J.R., BELL, J.G., BELL, M.V., HENDERSON, R.J. & TOCHER., D.J. (1993): The metabolism of phospholipids
and polyunsaturated fatty acids in fi sh. in: LAHLOU, B. & VITIELLO, P. (Eds) Aquaculture: Fundamental and
applied research. Coastal and estuarine studies 43. American Geophysical Union, Washington, DC, pp. 103–
124.
SARGENT, J.R., BELL, J.G., BELL, M.V., HENDERSON, R.J. & TOCHER, D.R. (1995): Requirement criteria for essential
fatty acids. Symposium of European Inland Fisheries Advisory Commission. J. appl. Ichthyol, 11, 183–198.
SARGENT, J., BELL, G., MCEVOY, L., TOCHER, D. & ESTEVEZ, A. (1999): Recent developments in the essential fatty acid
nutrition of fi sh. Aquaculture, 177, 191–199.
SIMON, J.A., HODGKINS, M.L., BROWNER, W.S., NEUHAUS, J.M., BERNERT, J.Y. & HULLEY, S.B. (1995): Serum fatty
acids and the risk of coronary heart disease. Am. J. Epidemiol., 42, 469–476.
VAN GELDER B.M., TIJHUIS M, KALMIJN S., KROMHOUT, D. (2007): Fish consumption, n–3 fatty acids, and subsequent
5-y cognitive decline in elderly men: the Zutphen Elderly Study. Am. J. clin. Nutr., 85, 1142–1147.