Comparative Biochemistry and Physiology, Part B 140 (2005) 437 – 443

www.elsevier.com/locate/cbpb

Proximate composition, fatty acid and lipid class composition of the

muscle from deep-sea teleosts and elasmobranchs
Hege M.W. akland, Iren S. Stoknes, Jannicke F. Remme, Margareth Kjerstad, Marianne Synnes*
Mb re Research, Box 5075, N-6021 Ålesund, Norway

Received 1 September 2004; received in revised form 11 November 2004; accepted 14 November 2004

Abstract

Proximate composition of muscle was determined for the following deep-sea fish species: roughhead grenadier (Macrourus berglax),
mora/deep-sea cod (Mora moro), Portuguese dogfish (Centroscymnus coelolepis), black dogfish (Centroscyllium fabricii), leafscale gulper
shark (Centrophorus squamosus), greater lantern shark (Etmopterus princeps), smalleyed rabbitfish/ghostshark (Hydrolagus affinis),
birdbeak dogfish (Deania calcea) and two species of smooth head (Alepocephalus bairdii and Alepocephalus agassizii). The first eight
species contained less than 1% fat in the muscle, while the last two contained 3.0% and 3.6% fat, respectively. Fatty acid and lipid class
composition was determined for the first five fish species and showed that the dominant class of lipids was phospholipids. The lipids
consisted mainly of polyunsaturated fatty acids (PUFA), and docosahexaenoic acid (DHA) was the dominant fatty acid. Roughhead grenadier
and mora showed resemblance to cod (Gadus morhua) regarding protein content, fat content and fatty acid composition. However, the
muscle from the deep-sea fish species did contain a higher proportion of arachidonic acid (20:4n-6) than cod muscle.
D 2004 Elsevier Inc. All rights reserved.

Keywords: Deep-sea fish species; Lipid content; Fatty acid composition; Polyunsaturated fatty acids; DHA; EPA; Arachidonic acid; Omega-3

1. Introduction flesh can be seen as bwhiteQ or bdarkQ meat (Suzuki, 1981).

When noncommercial fish species are being investigated
for the purpose of food consumption, research is primarily
focused on the fish fillets. Fish muscle is consistent of an
excellent amino acid composition and is a unique source for
nutrients and easily digestible protein (Venugopal et al.,
1996; Yanes et al., 1976). Its proteins and amino acid
profiles are quite similar to the muscle of terrestrial animals,
but the fish bodies are supported by a mass of water so the
muscle fibres require less structural support than the
muscles of land animals (Hultin, 1985). The fish muscle,
therefore, tends to have less connective tissue than muscle
from terrestrial animals, resulting in a more tender texture
(reviewed by Kristinsson and Rasco, 2000).
The protein composition in muscles varies by muscle
type, whether it is striated, smooth or cardiac muscle. The
striated muscles are the predominant form in fish, and the
* Corresponding author. Tel.: +47 70161366; fax: +47 70138978.
E-mail address: marianne@mfaa.no (M. Synnes).
1096-4959/$ - see front matter D 2004 Elsevier Inc. All rights reserved.
doi:10.1016/j.cbpc.2004.11.008
The white meat contains less lipids than the dark meat and is
usually composed of about 18% to 23% of protein,
depending on the species and time of harvesting (Suzuki,
1981). However, significant variations in protein content are
observed between fish species, and a relation seems to exist
between chemical composition and depth of occurrence
(Childress and Nygaard, 1973; Yancey et al., 1992).
The protein content and the water-holding capacity of the
proteins are also very important for the texture of the fish
flesh. Water-holding capacity refers to the ability of the
proteins to imbibe water and retain it against gravitational
force within a protein matrix. A high protein content and a
good water-holding capacity of proteins in fish muscle is
therefore important as it often improves the texture of the
fish flesh.
The nutritional importance of fish consumption is in
great extent associated with the content of omega-3 fatty
acids (n-3 FAs). Seafood is known as a rich source of the
n-3 FAs, and most marine oils are good sources of
eicosapentaenoic acid (EPA; 20:5n-3) and docosahexaenoic
438 H.M.W. a kland et al. / Comparative Biochemistry and Physiology, Part B 140 (2005) 437–443

acid (DHA; 22:6n-3). These fatty acids, particularly the Length and mass were determined for three individuals of
DHA, are known to play a major role in several biochemical each species used in the present study and are presented as
processes both in vivo and in vitro (Calder, 1997). DHA in meanFS.D.
the diet improves learning ability and the development of
the mammalian brain and has also a positive effect on 2.2. Analyses of moisture and ash
several diseases like hypertension, atherosclerosis and some
cancers (reviewed by Horrocks and Yeo, 1999). During the The muscle was homogenised in a food processor (Braun
last decades, there has been a focus on the n-3 fatty acids Combimax 600), and moisture content of 5 g of homoge-
and their positive effects. Despite of this, our western diet nised sample was determined by drying the sample in an
tends to have a too low content of n-3 fatty acids oven at 105 8C until a constant mass was obtained (AOAC,
(Kromhout, 2001). To counteract this, the increased 1990).
consumption of foods containing n-3 fatty acids should be Ash was determined by using the basic AOAC method
promoted. Fish has long been recognised as a good source (1990) heating the samples in the furnace at 550 8C for
of n-3 fatty acids. To identify the new exploited fish with 8–12 h.
high in n-3 fatty acids, the detailed information on their fatty The analyses were repeated three times, and the results
acids composition is required. are presented as meanFS.D. of determinations for triplicate
In this study, we investigated the proximate composition samples.
of muscle from 10 different deep-sea fish species and
examined further five of these species with a focus on their 2.3. Protein analyses
lipid class, fatty acid and amino acid composition.
Total protein was determined by the Kjeldahl method
(Ritzmann and Daniels, 1975) by which the concentration of
2. Materials and methods nitrogen is measured. A conversion factor of 6.25 was used
to convert total nitrogen to crude protein for all varieties of
2.1. Materials fish. The analyses were performed by The Norwegian Food
Control and Animal Welfare Authority, Dept. 2lesund,
Roughhead grenadier (Macrourus berglax), mora/deep- Norway.
sea cod (Mora moro), Portuguese dogfish (Centroscymnus The amino acid composition in fish muscle was
coelolepis), black dogfish (Centroscyllium fabricii), leaf- determined by HPLC, according to the method by Lindroth
scale gulper shark (Centrophorus squamosus), greater and Mopper (1979), at the Dept. of Biotechnology, NTNU,
lantern shark (Etmopterus princeps), smalleyed rabbitfish/ Trondheim, Norway. Hydrolysed solutions were diluted
ghostshark (Hydrolagus affinis), birdbeak dogfish (Deania 1:1000 and analysed by RP-HPLC with precolumn deriva-
calcea) and smooth head (Alepocephalus bairdii) were tisation with o-phthaldialdehyde and fluorescence detection.
obtained from Hatton Bank in the North Atlantic, while The samples were analysed on a Water NOVA PAK C-18,
Alepocephalus agassizii was caught outside of Greenland. 15 cm3.9 mm, 4 Am at flow 0.9 ml/min. This method does
All species were caught by long line at depths between not detect the amino acids cysteine, tryptophan and proline.
500 and 1600 m except for the smooth heads that were
caught by trawl. The fish were caught and stored at 20 2.4. Lipid analyses
8C until analysed. One hundred grams of flesh was cut out
from the same area of the dorsal muscle of three Total fat content, lipid fraction and fatty acid content of
individuals of each species after thawing. The flesh was the five first fish species mentioned above were obtained
used for all the analyses, and all the analyses were from SINTEF Fisheries and Aquaculture, Norway. The
performed at least three times. lipids were extracted using a modified method of Bligh and
Fish used in this study are presented below. Dyer (1959). Total fat content of the last five species was
determined gravimetrically after ethyl acetate extraction
Fish species Mass Length Capture depth according to Losnegard et al. (1979) by The Norwegian
(kg) (cm) (m) Food Control and Animal Welfare Authority, Dept.
Roughhead grenadier 3.7F0.2 78.5F3.5 1300–1600 2lesund, Norway. The lipid analyses were performed with
Mora/deep-sea cod 2.5F0.7 56.3F7.1 500–1200 two different methods because they were analysed at two
Portuguese dogfish 12.0F2.0 111.0F3.6 700–1600 different occasions.
Black dogfish 4.1F0.7 82.8F0.4 900–1600 Fatty acid methyl esters were prepared according to
Leafscale gulper shark 6.0F1.7 104.0F12.3 800–1600
Greater lantern shark 2.6F0.6 73.0F6.1 900–1600
Metcalf et al. (1966) and analysed on a Carbo Erba HRGC
Smalleyed rabbitfish 6.9F0.2 105.0F2.7 700–1500 5160 gas chromatograph equipped with an omegawax 320
Birdbeak dogfish 2.3F0.4 84.3F3.2 500–1500 (Supelco) glass capillary column, employing on-column
Smooth head (A. bairdii) 1.5F0.3 60.5F2.5 800–1200 injection and flame ionisation detector. Peaks were reported
Smooth head (A. agassizii) 3.1F0.5 65.7F6.8 800–1200 by a Shimadzu Chrompac C-3R computing integrator
 H.M.W. a kland et al. / Comparative Biochemistry and Physiology, Part B 140 (2005) 437–443 439

Table 1
Proximate composition of the muscle from various species of deep-sea fish
Fish species Fat Protein Water Ash
g/100 g wet weight
Roughhead grenadier (Macrourus berglax) 0.61F0.06 16.7F0.29 82.4F0.11 0.97F0.03
Mora/deep-sea cod (Mora moro) 0.41F0.03 17.6F0.11 81.6F0.04 1.04F0.03
Portuguese dogfish (Centroscymnus coelolepis) 0.95F0.07 20.8F0.21 79.9F0.17 0.93F0.02
Black dogfish (Centroscyllium fabricii) 0.70F0.02 17.1F0.51 84.1F0.09 0.84F0.04
Leafscale gulper shark (Centrophorus squamosus) 0.99F0.05 20.0F0.20 82.3F0.18 0.86F0.05
Greater lantern shark (Etmopterus princeps) 0.80F0.20 19.7F0.26 81.8F0.13 0.86F0.00
Smalleyed rabbitfish (Hydrolagus affinis) 0.95F0.10 18.3F0.61 81.6F0.37 1.14F0.02
Birdbeak dogfish (Deania calceus) 0.90F0.15 22.6F0.18 80.0F0.05 0.78F0.04
Smooth head (Alepocephalus bairdii) 3.00F0.04 10.1F0.18 87.2F0.03 0.78F0.04
Smooth head (Alepocephalus agassizii) 3.60F0.14 10.3F0.26 85.5F0.05 0.97F0.04
Means valuesFS.D. of determinations for triplicate samples.

identified by comparison with known standards (Nu-Chek
Prep, MN, USA) and quantified by means of the response
factor to the internal standard C21:0.
Lipid classes were separated by an Iatroscan thin layer
chromatography–flame ionization detector system (TLC–
FID). Chromarods SIII (Iatron Laboratories, Tokyo, Japan)
were first scanned twice through the Iatroscan FID
immediately before sample application to remove possible
contaminants from the rods. Chloroform solutions of the
extracts (1 ml, concentration 20 mg/ml) were spotted on the
Chromarods SIII by use of a single spotting action and a 10-
ml chromatographic syringe. After spotting, the rods were
conditioned in a constant humidity chamber for 8 min over a
saturated NaCl solution and then transferred immediately to
the developing tank. Two different types of solvent systems
were used: hexane/diethyl ether/formic acid (85:15:0.04) for
separating nonpolar lipids and chloroform/methanol/water
(70:35:3.5) for separating the polar lipids. The analyses
were performed three times.
3. Results
3.1. Proximate composition of the muscle from selected
deep-sea fish species
The content of fat, protein, moisture and ash was
examined in the muscle of the deep-sea fish species
roughhead grenadier, mora/deep-sea cod, Portuguese dog-
fish, black dogfish, leafscale gulper shark, greater lantern
shark, birdbeak dogfish, smalleyed rabbitfish/ghostshark
and smooth head. Most of the species examined showed a
similar distribution of fat, protein, moisture and ash (Table
1). The first eight species of fish contained less than 1% fat;
the protein content was between 16.7% and 22.6%, while
the moisture content varied from 79.9% to 84.1%. The
muscle of smooth head (A. bairdii and A. agassizii),
however, contained more fat than the others (3% and
3.6% fat, respectively) but contained only about 10%
protein and more than 85% moisture.
3.2. Distribution of lipid fractions in fish muscle of various
species of deep-sea fish
The lipid fractions in the muscle of roughhead grenadier,
mora, Portuguese dogfish, black dogfish and leafscale
gulper shark were further determined. The distribution of
cholesterol esters, triacylglycerols, free fatty acids, choles-
terol and phospholipids are presented as percent of total fat
in Table 2.
Based on these results, the fish species examined could
be divided into two groups according to the differences in
their lipid composition. Phospholipids were the dominant
class of lipids among all fish species and varied between
65% and 94% of total fat content. The muscle of the bony
species—roughhead grenadier and mora—contained more
free fatty acids and less phospholipid than the cartilaginous
species—Portuguese dogfish, black dogfish and leafscale
gulper shark (Table 2). The bony species contained more
cholesterol than the cartilaginous species, and roughhead

Table 2
Distribution of lipid fractions fish muscle of various species of deep-sea fish
Fish species Cholesterol esters Triacylgylcerols Free fatty acids Cholesterols Phospholipids
Percent of total fat content
Roughhead grenadier n.d. 5.48F0.18 20.08F2.94 9.46F0.59 64.99F3.71
Mora n.d. 0.48F0.05 17.1F0.13 7.82F0.06 74.61F0.02
Portuguese dogfish 1.87F0.04 0.99F0.04 1.77F0.16 4.84F0.04 87.54F0.11
Black dogfish n.d. n.d. 0.99F0.03 5.16F0.13 93.85F0.10
Leafscale gulper shark 0.77F0.10 0.88F0.21 1.68F0.16 4.46F0.90 86.83F1.74
Mean valuesFS.D. of determinations for triplicate samples; n.d.—not detected.
440 H.M.W. a kland et al. / Comparative Biochemistry and Physiology, Part B 140 (2005) 437–443

black dogfish and leafscale gulper shark is shown in Fig. 1.

Fig. 1. Distribution of saturated, monounsaturated and polyunsaturated fatty
acids in five species of deep-sea fish: roughhead grenadier, mora, Portuguese
dogfish, black dogfish and leafscale gulper shark, expressed as percent of
total fat content, meanFS.D. of determinations for triplicate samples.
grenadier also contained a significant amount of triacylgly-
cerols in contrast to the other species examined.
3.3. Fatty acid profiles of the fish muscle
The distribution of saturated (SFA), monounsaturated
(MUFA) and polyunsaturated fatty acids (PUFA) in the
muscle of roughhead grenadier, mora, Portuguese dogfish,
The results demonstrated that a significant part of the fatty
acids in the muscle was polyunsaturated fatty acids (PUFA).
The content of PUFA varied from 48% to 63% of total fat
content among the different species examined.
The fatty acid composition of the muscle of the above-
mentioned fish species is shown in Table 3. Among the
saturated fatty acids, the 16:0 fatty acid was the dominant
one. This was true for all the species examined. The fatty
acid 18:1 n-9 was the dominant monounsaturated fatty acid,
while 22:6 n-3 (DHA) was the dominant polyunsaturated
acid. Among the n-6 fatty acids, arachidonic acid (20:4n-6)
was the dominant fatty acid for all the species examined.
The n-3 FAs (DHA and EPA) accounted for between
85% and 88% of the PUFAs, of which DHA and EPA were
the most abundant. DHA accounted for 53% to 73% of the
PUFAs, and the muscles from elasmobranchs had the
highest content of DHA. Thus, the muscle of roughhead
grenadier and mora contained significantly more EPA than
the elasmobranchs examined.
As shown in Table 3, the n-3/n-6 ratio in muscle among
the different species was found to range between 5.5 and
7.5. The ratio of polyunsaturated versus saturated fatty acids
(PUFA/SFA) demonstrated a dominant percentage of poly-
unsaturated relative to saturated fatty acids.
The amount of n-3 FAs in the fish muscle from the
abovementioned species varied between 41% and 56%
(Table 4) when expressed as percent of total fat content.
However, the amount n-3 FAs was only between 0.2 and 0.5 g

Table 3
Fatty acid composition of muscle from various deep-sea fish species (%)
Fatty acids Roughhead grenadier Mora Portuguese dogfish Black dogfish Leafscale gulper shark
14:0 1.07F0.11 0.87F0.02 0.71F0.02 0.38F0.04 0.57F0.03
16:0 9.84F0.33 11.62F1.24 11.45F0.31 12.49F0.35 11.99F0.54
18:0 4.66F0.24 4.89F0.48 3.10F0.12 4.90F0.19 7.69F0.02
20:0 n.d. n.d. n.d. 0.13F0.03 n.d.
22:0 0.12F0.01 0.04F0.03 n.d. n.d. n.d.
16:1 3.44F0.23 2.49F0.09 4.41F0.04 0.72F0.04 0.85F0.04
18:1n-9 15.26F0.01 13.56F0.68 22.53F0.07 20.52F0.04 19.12F0.13
20:1 7.23F0.37 2.66F0.09 5.95F5.95 3.93F0.18 3.60F0.42
22:1 2.18F0.12 0.52F0.03 3.84F0.16 0.84F0.03 1.87F0.15
18:2n-6 1.10F0.00 1.06F0.00 0.99F0.01 1.15F0.04 0.8F0.21
18:3n-6 0.09F0.00 0.13F0.02 0.16F0.01 0.18F0.03 0.15F0.00
20:2n-6 0.28F0.03 0.15F0.01 0.18F0.01 0.19F0.01 0.21F0.05
20:3n-6 0.03F0.02 0.12F0.02 0.05F0.07 0.12F0.01 0.21F0.06
20:4n-6 5.73F0.26 6.00F0.89 5.88F0.13 6.97F0.28 6.24F0.05
18:3n-3 0.29F0.01 0.22F0.08 0.25F0.01 0.20F0.02 0.22F0.03
18:4n-3 0.43F0.01 0.17F0.01 0.10F0.01 0.27F0.10 0.29F0.05
20:4n-3 0.39F0.01 0.53F0.07 0.15F0.02 n.d. n.d.
20:5n-3 (EPA) 16.77F0.23 14.73F1.31 3.00F0.14 4.73F0.19 2.51F0.21
22:5n-3 1.54F0.05 3.01F00.49 4.11F0.16 3.62F0.09 4.00F0.00
22:6n-3 (DHA)
P 29.55F0.87 37.15F1.51 33.15F0.20 38.67F0.98 39.67F0.44
PSFA 15.69F0.03 17.41F0.69 15.26F0.21 17.89F0.06 20.26F0.60
MUFA 28.12F0.72 19.32F0.47 36.73F0.22 26.01F0.22 25.46F0.40
P
PUFA 56.20F0.74 63.27F1.16 48.01F0.01 56.10F0.28 54.29F0.20
n-3/n-6 6.8 7.4 5.6 5.5 6.1
PUFA/SFA 3.6 3.6 3.1 3.1 2.7
Mean valuesFS.D. of determination for triplicate samples. SFA—saturated fatty acids; PUFA—polyunsaturated fatty acids; DHA—docosahexaenoic acid;
EPA—eicosapentaenoic acid; n.d.—not detected.
 H.M.W. a kland et al. / Comparative Biochemistry and Physiology, Part B 140 (2005) 437–443
Table 4
Omega fatty acids expressed as percent of total fat content or as percent of total muscle weight (w/w)
Fish species Roughhead Mora
grenadier
n-3 as percent of total fat content 48.96 55.82
n-3 as percent of total muscle weight 0.30 0.23
Mean valuesFS.D. of determinations for triplicate samples.
a
These values were obtained from the National Institute of Nutrition and Seafood Research, Norway (Anonymous, 2002).
per 100 g fish muscle because of the low fat content in these
fish species.
3.4. Amino acid composition of fish muscle
The amino acid composition of the muscle from the
above mentioned species was determined, and the profile
was similar for all the species here represented by the amino
acid composition of the muscle from roughhead grenadier
(Fig. 2). The dominant amino acid was lysine, but relatively
high amounts of asparagine, glutamine, arginine and leucine
were observed. It was, however, difficult to discriminate
between glycine and arginine in our amino acid analyses.
The amount of the amino acids cysteine, tryptophan and
proline was not determined, but these amino acids would
only account for 5% of total protein since the amino acid
composition of 95% of total protein was determined. The
three missing amino acids would therefore not change the
amino acid profile significantly.
4. Discussion
During the last few years, there has been an increasing
investigation of deep-sea fish species, with the intention of
exploring so far underutilised fish species that can replace
or supplement the available commercial species. It is
therefore important that the new species can compete with
Fig. 2. Amino acid composition (as percent total protein) of the muscle of
roughhead grenadier, a representative of deep-sea fish species.
441
Portuguese Black Leafscale Cod Piked
dogfish dogfish gulper shark dogfish
40.75 47.49 46.68 54.3a 31.4a
0.39 0.33 0.46 0.16a 2.01a
the traditional species regarding both taste and nutritional
values. In this study, we determined the chemical
composition of 10 deep-sea species, roughhead grenadier,
mora/deep-sea cod, Portuguese dogfish, black dogfish,
leafscale gulper shark, greater lantern shark, birdbeak
dogfish, smalleyed rabbitfish/ghostshark and two species
of smooth head.
The data on fat content and fatty acid composition are
important when utilisation of new species of fish is
considered. This is because fish is considered to be not
only food with good source of quality protein but also food
with healthy components. Fish can be grouped into four
categories according to their fat content: lean fish (b2%),
low fat (2–4%), medium fat (4–8%) and high fat (N8%;
Ackman, 1989). Of the 10 fish species investigated, the
eight first were considered to be lean fish, having fat content
less than 1% (Table 1), while the last two species were
considered to be low fat fish.
The protein content is important when considering
quality and texture of the fish muscle. Fish muscles that
contain small amounts of protein tend to loose much water
upon cooking, which ruins the texture of the meat. The
present study showed that, with respect to their protein
content (Table 1), roughhead grenadier and mora may be
compared to the well-known cod (Gadus morhua), which
has a protein content of 18.1% (Anonymous, 2002). The
cartilaginous species examined also seemed to have a high
protein content (Table 1) and could be compared to their
better known congener, piked dogfish, which has a protein
content of 17.9% (Anonymous, 2002). The two species of
smooth head differed from the other species examined in
the present study, with much lower protein content. Such
low protein content have, however, also been observed for
several other fish species like blue whiting (9.8%; Brennan
and Gormley, 1999) and northern wolffish (10.5%;
unpublished results).
In a previous study, Brennan and Gormley (1999)
determined the protein content of some of the species
examined in the present study: birdbeak dogfish (25.2%),
greater lantern shark (21.5%), leafscale gulper shark
(22.0%), mora (17.3%), Portuguese dogfish (22.4%) and
Baird’s smooth head (12.9%). In another study, Gormley et
al. (1994) predicted the protein content of Baird’s smooth
head to be only 6.4%. Their results seem to be slightly
higher than our results regarding the protein content of
elasmobranchs. The protein content of mora is, however, the
same as predicted in the present study.
442 H.M.W. a kland et al. / Comparative Biochemistry and Physiology, Part B 140 (2005) 437–443
Variations in determined protein content may be a result
of several factors. When determining the protein content by
the Kjeldahl method, a conversion factor of 6.25 is used to
convert total nitrogen to crude protein. It should however be
noted that a high content of nonprotein nitrogen (NPN) will
give an overestimate of the actual protein content. Elasmo-
branchs have a high content of NPN in the form of
trimethylamine oxide (TMAO) and urea (reviewed in
Yancey et al., 2002; Hazon et al., 2003).
Oehlenschläger (1992) proposed that a factor of 5.4
would be suitable to estimate protein content from total
measured nitrogen in elasmobranchs species. However, for
deep-sea species, the capture depth may be important since
it has been demonstrated that amount of TMAO in the
muscle increases with increasing depth (Kelly and Yancey,
1999). Deep-sea elasmobranchs have less urea and more
TMAO, making the 5.4 correction possibly wrong. Deep-
sea teleosts also have a lot of NPN in the form of TMAO
(Gillett et al., 1997; Kelly and Yancey, 1999), making the
estimated protein content in these species probably too
high as well. Depth of occurrence of the examined fish
species might therefore interfere with the protein measure-
ments, and as a result of this, the protein values reported
in Table 1 may be overestimated.
In our further investigations, we decided to evaluate the
lipid part of the deep-sea species roughhead grenadier,
mora, Portuguese dogfish, black dogfish and leafscale
gulper shark since they seem to be of greatest commercial
interest. The first two are bony fish species, while the last
three are cartilaginous fish species.
The dominant class of lipids among all the species
examined was the phospholipids. However, a difference
between the bony fish species and the cartilaginous species
examined could be observed. The bony species had a
significantly higher content of free fatty acids and also a
higher content of cholesterol than the cartilaginous species
(Table 2). A high content of free fatty acids may be due to
degradation of the lipids. In the present study, the bony and
the cartilaginous species were treated similarly after capture
so it is not likely that the higher level of free fatty acids
observed in muscle from roughhead grenadier and mora was
due to degradation of the lipids caused by differences in
storage of the fish.
The lipids of the muscle from both bony and cartilagi-
nous fish species examined in the present study consisted
mainly of polyunsaturated fatty acids (PUFA; Fig. 1). The n-
3 FAs and n-6 FAs are two biochemical families within the
PUFAs, and they also have different biological effects
(James and Cleland, 1996). Traditionally, fish with high fat
content, like rainbow trout, has been considered to be
nutritional important species since they have a relatively
high content of n-3 FAs. However, it has been demonstrated
that there is an inverse relationship between amount of n-3
FAs and total fat content (Kristinsson and Rasco, 2000).
This implies that it is important from the nutritional point of
view to place attention upon getting species with high
proportions of n-3 FAs instead of only focusing on the fat
content. Previous studies have also demonstrated a benefi-
cial role of lean fish, as well as fatty fish consumption, in the
prevention of cardiovascular diseases (Kromhout, 2001).
The fish species from the present study cannot be
compared to fat fish species when it comes to total amount
of n-3 FAs. However, roughhead grenadier and mora can
be compared to cod when it comes to total fat content and
content of n-3 fatty acids (Tables 1 and 4). Cod contains
only 0.3% fat of which 54.3% is n-3 fatty acids
(Anonymous, 2002).
Compared to piked dogfish, the deep-sea dogfishes more
commercial congener, we observed that Portuguese dogfish,
black dogfish and leafscale gulper shark were much leaner.
Piked dogfish is considered to be a good source of n-3 FAs
with a fat content in muscle of about 6% and 31% of the
total fat content being n-3 FAs (Anonymous, 2002). Thus,
although the proportion of n-3 FAs are higher in Portuguese
dogfish, black dogfish and leafscale gulper shark than in
piked dogfish (Table 4), the amount of n-3 FAs per 100 g
fish muscle is much lower. In marine fish, the PUFA
comprise mainly the n-3 fatty acids eicosapentaenoic acid
(EPA) and docosahexaenoic acid (DHA; Ackman, 1989).
However, the ratio of DHA versus EPA varies among
species or among individuals. Thus, fish species with a high
content of n-3 FA in the muscle is not necessarily a good
source of DHA.
In the present study, we observed that DHA and EPA
accounted for 36–52% of the total fatty acids in the muscle
of the species examined (Table 3). The bony species,
roughhead grenadier and mora contained almost five times
more EPA than the cartilaginous species. DHA accounted
alone for about 30–40% of total fat content, which is
considered to be particularly high. However, it has
previously been demonstrated such high ratio of DHA in
the muscle of the tuna Euthynnus (Katsuwonus) pelamis
(Saito et al., 1997).
Arachidonic acid (20:4n-6) was the dominant n-6 fatty
acid in muscle from all the deep-sea fish species examined
(Table 3). The content of arachidonic acid was more than
twice as high as the level found in muscle from their
brelativesQ living in the upper strata, cod and piked dogfish
(1.8% and 2.8%, Anonymous, 2002).
Piggott and Tucker (1990) suggested that the n-3/n-6
ratio is a better index in comparing relative nutritional value
of fish oils of different species. A n-3/n-6 ratio of 1:1 is
considered to be optimal for nutritional purposes (Simo-
poulos, 1989), particularly when n-3 FAs consist mainly of
EPA and DHA. Today, this balance in the usual diet has
been severely shifted, and the amount of n-3 FAs in the diet
has decreased, while the amount of n-6 FAs has increased. A
dietary intake of fish with high ratio of n-3/n-6 would
therefore be beneficial.
The n-3/n-6 ratio was found to range between 5.5 and 7.5
among the fish species in the present study (Table 3). These
results indicate that the deep-sea fish species from the
 H.M.W. a kland et al. / Comparative Biochemistry and Physiology, Part B 140 (2005) 437–443
present study contains a particularly good lipid composition
for nutritional purposes. It should however be noted that the
net content of n-3 and n-6 is low because of the low lipid
content of these fish species, and only regular consumption
of such fish species could contribute to increase the amount
of n-3 and n-6 fatty acids in the diet.
Fish muscle is known to contain an excellent amino acid
composition (Venugopal et al., 1996; Yanes et al., 1976) and
is a unique source of physiological beneficial amino acids
(arginine, histidine, lysine, taurine; Marshall, 1994; Metzner
et al., 2001). Our studies indicated that these fish species
were particularly rich in lysine (Fig. 2), but otherwise, the
amino acid profiles look quite similar to what has been
published for several other fish species (Anonymous, 2002).
This study demonstrates that several of the underutilised
deep-sea species are able to compete with more commer-
cially utilised species in terms of nutritional value, and they
can definitely also compete when it comes to taste
(unpublished work).
Acknowledgements
Andreas Wammer, Ann H. Hellevik, Jan Erich
Rbnneberg, Jannicke Martinussen and Wenche Emblem,
all from Mbre Research, are gratefully acknowledged for
skillfully performing preparative and analytical laboratory
work. This study was supported by the Research Council
of Norway.
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