Journal of Food Composition and Analysis 23 (2010) 716–725

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Journal of Food Composition and Analysis

journal homepage: www.elsevier.com/locate/jfca

Original Article

Chemical composition, cholesterol, fatty acid and amino acid in two populations
of brown crab Cancer pagurus: Ecological and human health implications
Sara Barrento a, António Marques a,*, Bárbara Teixeira a, Rogério Mendes a, Narcisa Bandarra a,
Paulo Vaz-Pires b,c, Maria Leonor Nunes a
a
Research Unit of Upgrading of Fishery and Farmed Products (U-VPPA), National Institute of Biological Resources (INRB I.P./L-IPIMAR), Avenida de Brası´lia, 1449-006 Lisboa, Portugal
b
Institute of Biomedical Sciences Abel Salazar (ICBAS-UP), University of Porto, Largo Professor Abel Salazar 2, 4099-003 Porto, Portugal
c
Centre of Marine and Environmental Research, University of Porto (CIIMAR-UP), Rua dos Bragas 289, 4050-123 Porto, Portugal

A R T I C L E I N F O A B S T R A C T

Article history: The brown crab, Cancer pagurus, is a valued decapod species captured mainly in the UK and France. In
Received 8 July 2009 Portugal, Scottish crabs and females of all species are less expensive than crabs caught in the English
Received in revised form 28 January 2010 Channel and males. In this work the proximate chemical composition, cholesterol, fatty acid and amino
Accepted 16 March 2010
acid content of female and male C. pagurus edible tissues from the Scottish coast and English Channel
were compared. There was no evidence that the fishing ground influenced the chemical composition of
Keywords: tissues, but there were significant differences between tissues and sexes. Muscle was richer in protein,
Crustaceans
but poorer in fat and cholesterol, than gonads and hepatopancreas. Ovaries had more protein, fat,
Cancer pagurus
cholesterol and amino acids than testes. The fatty acid profiles in muscle and gonads were dominated by
Hepatopancreas
Gonads PUFA, while hepatopancreas was richer in MUFA and SFA. Lower n-3 fatty acid content and n-3/n-6 fatty
Muscle acid ratio in hepatopancreas contributed to higher atherogenic and thrombogenic indices. Considering
Cholesterol the chemical composition, there is no reason for price differentiation between crabs from different
Fatty acids locations. As far as sex is concerned the principal difference that might increase male crabs’ value is the
Amino acids meat yield content of claws, which was higher than females’ claws.
Nutritional quality ß 2010 Elsevier Inc. All rights reserved.
Analysis of fishing ground and nutrition
Biodiversity
Food analysis
Food composition

1. Introduction meat (gonads and hepatopancreas) are appreciated. In these

Worldwide, crustaceans are highly appreciated and are consid-
ered luxury seafood items. Although their frequent consumption is
not advisable in general, either because of allergenic reactions or the
supposedly high cholesterol content, there are a growing number of
studies promoting crustacean consumption (Rosa and Nunes,
2003a; Gökoolu and Yerlikaya, 2003; Küçükgülmez et al., 2006;
Chen et al., 2007; Teixeira et al., 2008; Barrento et al., 2008a,
2009a,b).
Despite this controversy, crustaceans are widely appreciated and
consumed in Mediterranean countries. In particular, the crab Cancer
pagurus is a valued decapod species in France, Spain, Portugal and
Italy, where the muscle from claws and abdomen and the brown
* Corresponding author. Tel.: +351 21 3027025; fax: +351 21 3015948.
E-mail addresses: amarques@ipimar.pt, marques_am@yahoo.com (A. Marques).
URL: http://www.inrb.pt/ipimar/investigacao/unidade-de-investigacao-de-
valorizacao-dos-produtos-da-pesca-e-da-aquicultura-
countries the market trend is still the commercialization of live
crabs harvested in the waters off Britain (31,079 tonnes), Ireland
(11,525 tonnes) and France (5724 tonnes) (Tully et al., 2006; data
from 2007, EUROSTAT, 2009). The major export countries to Portugal
are UK and France. This species shows different market prices
according to the fishing ground and sex. In fact, crabs caught off the
English Channel are usually more expensive than those caught off
Welsh, Scottish and Irish coasts, presumably due to the intrinsic
quality of these populations (e.g. bigger animals that undergo fewer
hours of transport and which are therefore more vigorous upon
arrival) (Barrento et al., 2008b). In addition, males are more
expensive than females (Brown and Bennett, 1980).
It is well known that the biochemical composition of edible
tissues of marine invertebrates is influenced by their nutritional
habits, age, sex, season, seawater temperature and salinity
(Chapelle, 1977; Oliveira et al., 2007; Souchet and Laplante,
2007). Since crab populations from different fishing grounds might
have distinct biochemical patterns in the edible tissues, the
purpose of this study was to determine the nutritional quality of

0889-1575/$ – see front matter ß 2010 Elsevier Inc. All rights reserved.
doi:10.1016/j.jfca.2010.03.019
 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725 717

Table 1
Summary of reference material elemental concentration (mg g1 DW; n = 4) and detection limits (DL) of certified reference material (standard deviation) analyzed by the
different techniques.

Elements Technique DL Certified biological material Certified Present work

Moisture Drying nd Canned matrix meat – SMRD 2000 68.8  0.1 68.7  0.0
Protein Kjeldahl nd Canned matrix meat – SMRD 2000 1.63  0.05 1.62  0.06
Total lipids Soxhlet 0.01 Canned matrix meat – SMRD 2000 14.3  0.4 14.4  0.1
Ash Combustion 0.1 Canned matrix meat – SMRD 2000 2.65  0.07 2.69  0.03
Fatty acids
14:0 GC 0.4-1 Beef pork fat blend – BCR-163 2.29  0.04 2.23  0.04
16:0 GC 0.4-1 Beef pork fat blend – BCR-163 25.96  0.30 25.39  0.017
16:1 GC 0.4-1 Beef pork fat blend – BCR-163 2.58  0.16 2.22  0.03a
18:0 GC 0.4-1 Beef pork fat blend – BCR-163 18.29  0.16 17.65  0.06
18:1 GC 0.4-1 Beef pork fat blend – BCR-163 38.34  0.36 38.65  0.16b
18:2 GC 0.4-1 Beef pork fat blend – BCR-163 7.05  0.17 7.19  0.03c
18:3 GC 0.4-1 Beef pork fat blend – BCR-163 0.86  0.14 0.81  0.01d
Amino acids HPLC nd No reference material used nd nd
Cholesterol GC nd No reference material used nd nd

Reference materials suppliers and location: SMRD-2000 (Swedish Meats R&D and Scan Foods/National Food Administration, Sweden), BCR-163 (Institute for Reference
Materials and Measurement, Belgium). Abbreviations: gas chromatography (GC), high-performance liquid chromatography (HPLC), not determined (nd).
a
16:1n-7 + 16:1n-5.
b
18:1n-9 + 18:1n-7.
c
18:2n-6.
d
8:3n-3.

C. pagurus harvested in the English Channel and Scottish coast by
determining the proximate chemical composition, amino acid and
fatty acid profiles, and cholesterol content. Data were also
analyzed to evaluate the nutritional quality of C. pagurus edible
tissues to human consumption by calculating the atherogenic and
thrombogenic indices, the essential amino acid scores and three
ratios related with fatty acid content: DHA/EPA (docosahexaenoic
acid/eicosapentaenoic acid), EPA/DHA and PUFA/SFA (polyunsatu-
rated fatty acids/saturated fatty acids).
2. Materials and methods
2.1. Biological material
Twenty C. pagurus specimens caught off the Scottish coast in
summer (SC; 10 females and 10 males) and 20 specimens from the
English Channel (EC; 10 females and 10 males) were purchased live
and transported to the laboratory. The biological material was the
same used in previous studies published by Barrento et al.
(2009a,c). Animals were kept under refrigerated conditions (5 8C)
during 1 h to decrease their metabolism, and stunned before being
euthanized by piercing the two nerve centres by means of a
stainless steel rod. The rod was inserted through one of the eyes
and through the vent as recommended by the Codex Alimentarius
Commission (FAO/WHO, 1983).
Several parameters were recorded for each crab: sex, total
weight (Omega PP 50\1; d = 1 g), claw muscle, gonad and
hepatopancreas weight (Mettler PM 400; d = 0.001 g), carapace
width and length (digital caliper Comecta; 0.01 mm), and ovarian
maturity stage. The ovarian maturity scale was adapted from
Edwards (1979). Females were classified as immature/no ovary
development (Stage 1: S1); initial ovary development (Stage 2: S2);
developing orange ovaries extending into carapace (Stage 3: S3);
and ripe, carapace full of bright red ovary material (Stage 4: S4).
Gonadosomatic (GSI) and hepatosomatic (HI) indices were
calculated as well as claws muscle meat yield (MY) and the total
meat (TMY) yield. The GSI, HI and MY, were calculated as follows:
(tissue wet weight/body wet weight)  100; the TMY was
calculated according to the formula: (gonad wet weight + hepa-
topancreas wet weight + claw muscle wet weight)/(body wet
weight)  100. For each tissue the edible content was also
calculated as follows: (tissue wet weight/sum of all edible tissues
wet weight)  100. For the biochemical analyses, the edible tissues
of at least two crabs were pooled taking into account sex, ovarian
maturity stage and location, and were homogenized with a grinder
(Retsch Grindomix GM200, Düsseldorf, Germany; 5000 rpm;
material: PP cup and stainless steel knifes), vacuum packed and
stored at 20 8C. A portion of each frozen sample was freeze-dried
for 48 h at 50 8C and low pressure (approximately 101 atm; Heto
power dry LL 3000), and stored at 20 8C under controlled
moisture conditions (vacuum packed) until further analyses.
2.2. Proximate chemical composition and energy content
Moisture, ash, protein and lipid contents were determined in
duplicate for each sample, according to the AOAC (2005) methods
that were validated with reference material (Table 1). Briefly, the
moisture content was obtained by drying the sample overnight at
105 8C (laboratory heater, P-Selecta 207); ash was quantified after
combustion for 16 h at 550 8C (muffle furnace, Heraeus Hanau,
TYPMR170); crude protein content was determined by the Kjeldahl
method using an automatic distillation and titration unit (VELP
Scientifica, UDK152) with a conversion factor of 6.25; and total
lipid content was determined with the Soxhlet extraction method
using ethyl ether (40–60 8C; 7 h; heater plate SBS Instruments
PC6L). The results were expressed in g per 100 g wet weight. The
energy content was estimated as: proteins, 4.27 kcal g1 wet
weight; lipids, 9.02 kcal g1 wet weight; 1 kcal = 4.184 kJ (FAO,
1987).
2.3. Cholesterol
Cholesterol content was determined in duplicate for each
sample, and was based on the modified procedure of Naeemi et al.
(1995). Briefly, each sample (300 mg of dry weight) was combined
with 260 mL of the internal standard solution (5a-cholestane
Sigma; 5 mg mL1 cyclohexane, Merck), 3 mL of saturated
methanolic potassium hydroxide solution (2 M, Merck), and
3 mL of methanol pro analysis (Merck). Following heating
(80 8C; 30 min water bath), samples were cooled and supplied
with 250 mL of magnesium chloride solution (1 M, Merck) and
1.5 mL of cyclohexane pro analysis (Merck). Samples were shaken
and centrifuged (1500  g; 4 min, centrifuge Sigma 2K15) until
phase separation. The moisture content of the upper phase was
removed with anhydrous sodium sulfate (Panreac). The cholesterol
in the upper phase (2 mL) was separated by gas chromatography
718 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725
(Varian Star 3400 Cx) using helium as carrier gas at a flow rate of
1 mL min1 in a flame ionization detector and a fused silica
capillary CP-Sil 8 CB column (30 m length  0.25 mm internal
diameter, 0.25 mm film thickness; J&W Scientific). The tempera-
tures of the oven, injector, and detector were 280, 285, and 300 8C,
respectively. Cholesterol was identified and quantified by com-
parison with the calibration curve of a pure cholesterol standard
with 5a-cholestane (Sigma). Cholesterol/cholestane peak area
ratios obtained with the Varian software were plotted with
cholesterol concentrations, and a straight line was fitted to data
points by linear regression.
2.4. Fatty acid analysis
Fatty acid methyl esters (FAME) profile from nonpolar and polar
lipids was determined in duplicate for each sample, and was based
on the experimental procedure of Cohen et al. (1988). Each sample
(300 mg of dry weight) was dissolved in 5 mL of acetyl chloride/
methanol (1:19 v/v; Merck), shaken, and heated (80 8C; 1 h). After
cooling, 1 mL of Milli-Q distilled water and 2 mL of n-heptane pro
analysis (Merck) were added, and samples were shaken and
centrifuged (2000  g; 5 min, Sigma 2k15) until phase separation.
The moisture content of the upper phase was removed with
anhydrous sodium sulfate (Panreac). An aliquot (2 mL) of the upper
phase was then injected onto a gas chromatograph (Varian Star
3800 Cp) equipped with an autosampler and fitted with a flame
ionization detector at 250 8C for FAME analysis. The separation was
carried out with helium as carrier gas at a flow rate of 1 mL min1,
in a capillary column DB-WAX (30 m length  0.32 mm internal
diameter; 0.25 mm film thickness; Hewlett–Packard) programmed
at 180 8C for 5 min, raised to 220 at 4 8C min1, and maintained at
220 8C for 25 min, with the injector at 250 8C. FAME were identified
by comparing retention times with those of Sigma standards.
Quantitative data were calculated using the peak area ratio
(percent of total fatty acids) and the Varian software. The fatty acid
determination method was validated with reference material
(Table 1).
2.5. Amino acids
Amino acid concentration was determined in duplicate for
each sample. To extract total amino acids (protein bound + free),
40–90 mg of sample was placed in 10 mL ampoules with 3 mL of
6 M HCl (Merck) with 0.1% phenol (Merck), according to the
method described by AOAC (2005). Ampules were vacuum-
sealed, and samples were hydrolyzed at 110 8C for 24 h;
hydrolysates were filtered (0.45 mm pore size, Life sciences)
and dissolved with Milli-Q distilled water to 20 mL. Samples were
frozen at 80 8C and freeze-dried during 48 h at 50 8C under low
pressure (approximately 101 atm). Samples were dissolved in
5 mL of 0.1 M HCl (Merck) and stored at 80 8C until amino acid
separation. After thawing samples were filtered (0.2 mm pore size
Life sciences) and separation was performed with high-perform-
ance liquid chromatography (Agilent 1100 HPLC) using pre-
column o-phthalaldehyde and 3-mercaptopropionic acid in
borate buffer (OPA, Agilent Technologies) and 9-fluorenylmethyl-
chloroformate in acetonitrile (FMOC; Agilent Technologies)
derivatization, a Phenomenex Gemini ODS C18 guard column
(4 mm  3 mm), and a Phenomenex Gemini ODS C18 110A
column (4.6 mm  150 mm, 5 mm). The solvents and gradient
conditions were those described by Henderson et al. (2000).
Detection wavelengths were set at UV 338 and 262 nm and
fluorescence 340/450 and 266/305 nm. The identity and quantity
of the amino acids were assessed by comparison with the
retention times and peak areas of standard amino acids (Sigma–
Aldrich) using norvaline as internal standard.
2.6. Nutritional quality
The propensity of crab’s tissue to promote the incidence of
coronary heart disease, atherogenic (AI) and thrombogenic (TI)
indices were calculated by using the Ulbricht and Southgate (1991)
equations (see Eq. (1)).
½12 : 0 þ ð4  14 : 0Þ þ 16 : 0
AI ¼ P P ; (1)
½ MUFA þ PUFAðn  6Þ þ ðn  3Þ
ð14 : 0 þ 16 : 0 þ 18 : 0Þ
TI ¼ P P
½ð0:5  MUFAÞ þ ð0:5  PUFAðn  6ÞÞ
P
þ ð3  PUFAðn  3Þ þ ððn  3Þ=ðn  6ÞÞ
where MUFA means monounsaturated fatty acids and PUFA means
polyunsaturated fatty acids.
Three ratios related with fatty acid content were also
calculated: DHA/EPA (docosahexaenoic acid/eicosapentaenoic
acid), EPA/DHA and PUFA/SFA (saturated fatty acids), in order to
allow comparisons with the United Kingdom Department of Health
recommendations (HMSO, 1994).
Essential amino acid scores (AS) were also calculated
with respect to reference amino acid requirements for adults:
isoleucine (30 mg g1 protein), leucine (59 mg g1 protein), lysine
(45 mg g1 protein), methionine (16 mg g1 protein), phenylala-
nine + tyrosine (38 mg g1 protein), threonine (23 mg g1
protein), valine (39 mg g1 protein) and histidine (15 mg g1 pro-
protein) (FAO/WHO/UNU, 2007). This was achieved by comparing
the content of each essential amino acid in the protein/diet with its
requirements, according to the equation: AS (%) = (mg of
amino acid per g of protein/mg of amino acid requirement)  100.
The ratio of essential amino acids to non-essential amino acids was
also calculated.
2.7. Statistics
Main differences among tissues, sexes and locations were
tested with analysis of variance (ANOVA). Whenever necessary,
data were transformed to satisfy normal distribution and
homoscedasticity requirements, followed by nonparametric anal-
ysis of variance (Kruskall–Wallis), if transformed data could not
meet these assumptions. The logarithmic transformation was used
whenever the standard deviation was proportional to the means
(i.e. there is a constant coefficient of variation), whereas the square
root transformation was used when the group variances were
proportional to the means, that is when the variances increase as
the means increase (Zar, 1999). Principal component analysis
(PCA) was also employed to reduce the multidimensional data sets
of several elements to lower dimensions, thus simplifying the
presentation and interpretation of data. All statistical analyses
were tested at 0.05 level of probability with the software
STATISTICATM 6.1. (Statsoft, Inc., Tulsa, OK 74104, USA).
3. Results
3.1. Proximate chemical composition
The biometric data and proximate chemical composition of
brown crab are presented in Tables 2 and 3. Muscle and
hepatopancreas were responsible by the major edible contribution.
Muscle represented the highest content in male crabs (50.7–52.5%)
independently of location compared to females (38.3–42.4%),
whereas the contribution of gonads from both locations was higher
in females (14.2–15.4%) than in males (4.6–6.0%). The content of
hepatopancreas (43.3–46.3%) was constant between sexes
and locations. Regarding the proximate chemical composition
 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725 719

Table 2 (p < 0.01) than those of the EC, independently of sex. The
Biometric data of C. pagurus from both locations and sexes (average  standard
proportion of n-3, the ratios n-3/n-6, PUFA/SFA and IT were
deviation).
constant in respect to sex and location (Table 4). In contrast,
SF EF SM EM hepatopancreas revealed few differences related either with sex or
Weight (g) 748  25 751  27 829  155 977  104 location as only 22:5n-3 had a lower proportion in females than in
CL (mm) 107.38  1.77 109.93  5.88 98.44  3.71 110.16  3.71 males and 16:1n-5 had a higher proportion in crabs from EC and
CW (mm) 163.32  2.26 168.75  9.41 156.30  5.93 177.20  5.96 among these, females had more 16:1n-5 than males. Most
TMY (%) 22.5  3.8 23.5  2.2 22.0  9.9 23.8  3.6
differences found in gonads were related to sex, as 16:1n-7,
Abbreviations: SF, females from the Scottish coast; EF, females from the English 18:1n-9, 22:5n-3 and, the ratio n-3 to n-6 were all proportionally
Channel; SM, males from the Scottish coast; EM, males from the English Channel;
higher in females than in males. On the other hand 18:0, 16:3n-3
CL, carapace length; CW, carapace width; TMY, total meat yield. For each location
and sex n = 10. and 20:5n-3 were proportionally higher in males. The only
population differences were observed in SC crabs which had
proportionally more 20:1n-9. The fatty acid profiles were
(Table 3), significant differences were detected among tissues statistically different between tissues (Table 5). Muscle and
(Table 3). Few sex and location differences were observed in the gonads had a similar pattern dominated by PUFA, followed by
proximate chemical composition of muscle and hepatopancreas. MUFA and SFA, while hepatopancreas had proportionally more
On the contrary, gonads chemical composition showed marked MUFA and SFA. Hepatopancreas had proportionally more 14:0,
differences between sexes independently of location, as females 15:0, 16:1n-7, 20:1n-9, and 20:2n-6 than muscle and gonads, but
had more proteins, fat, cholesterol and energy than males but less less 16:1n-5, 20:4n-6, and 20:5n-3 (p < 0.01). The proportion of n-
ash and moisture. Muscle and ovaries had high protein content 3, n-6 and the ratio n-3/n-6 was lower in hepatopancreas than in
compared to hepatopancreas. Muscle also had the lowest fat muscle and gonads, whereas IA and IT were higher.
content and cholesterol concentration compared to gonads and
hepatopancreas. Concerning the female maturation stage, crabs 3.3. Amino acids
from both locations were mainly in the S3 stage (SC: 40%; EC: 50%),
while the remaining specimens were S2 (SC: 30%; EC: 25%) and S4 Non-essential amino acids (NEAA) dominated the protein
(SC: 30%; EC: 25%). content in all tissues of crabs from both locations and sexes,
namely glutamic acid, aspartic acid and glycine (Table 6). As to
3.2. Fatty acids essential amino acids (EAA), all tissues had mostly arginine, leucine
and threonine. Generally, amino acid concentration in the edible
In all tissues the main SFA, MUFA and PUFA were respectively tissues showed few differences in respect to population and more
palmitic acid (16:0), oleic acid (18:1n-9) and, eicosapentaenoic in relation to sex. In muscle, males had more methionine and
(EPA; 20:5n-3) and docosahexaenoic acids (DHA; 22:6n-3) (Table taurine than females. On the other hand, SM crabs had higher
4). The fatty acids profile of muscle showed differences mainly concentration of aminoacids like valine, leucine, asparagine and
according to location, as crabs from the SC had higher proportion of serine. In hepatopancreas most differences were related to sex, in
14:0, 18:1n-9 and 18:2n-6, and lower 18:1n-7 and 16:4n-3 which male crabs had usually higher concentration of several

Table 3
Proximate chemical composition (%), cholesterol content (mg 100 g1 wet weight) and energy content (kcal 100 g1 tissue wet weight) in edible tissues of Cancer pagurus
(average  standard deviation).

SF EF SM EM p

Muscle
MY 8.5  1.0a 9.1  3.6a 11.9  1.1a 12.4  2.1a <0.001
EC 38.3  5.2b 42.4  5.4b 50.7  5.0a 52.5  5.2a <0.001
Moisture 77.8  2.1a 77.4  1.9a 74.6  2.2b 77.3  1.3a 0.001
Ash 2.1  0.2a 2.2  0.2a 1.9  0.1b 2.2  0.2a 0.003
Proteins 16.4  4.2b 18.8  1.3ab 20.5  1.7a 17.9  0.8b 0.001
Fat 0.3  0.1a 0.4  0.1a 0.2  0.1a 0.4  0.1a 0.050
Cholesterol 37.3  4.7a 40.6  4.7a 37.0  9.1a 40.7  3.6a 0.233
Energy 85.9  10.1a 86.7  7.6a 95.4  10.3a 87.9  4.8a 0.324
Hepatopancreas
HI 10.4  1.8a 9.3  3.4a 10.2  2.2a 10.2  2.3a 0.806
EC 46.3  4.5a 43.3  4.7a 43.3  6.2a 42.9  6.3a 0.503
Moisture 67.6  6.7a 69.8  5.6a 61.8  7.3ab 61.1  2.3b 0.006
Ash 3.0  0.4b 3.1  0.9b 5.0  1.2a 6.6  0.6a 0.006
Proteins 12.2  1.6ab 9.4  4.6b 13.3  1.9ab 14.3  1.8a <0.001
Fat 10.2  3.6a 10.0  1.6a 12.2  3.4a 10.8  2.3a 0.056
Cholesterol 116.6  27.3a 115.6  57.1a 112.4  10.1a 126.7  13.0a 0.518
Energy 167.5  52.7a 154.9  24.9a 195.7  46.9a 184.5  15.3a 0.103
Gonads
GSI 3.6  2.0a 3.6  3.1a 1.4  0.7b 1.1  0.5b <0.001
EC 15.4  6.4a 14.2  6.0a 6.0  3.2b 4.6  1.6b <0.001
Moisture 56.8  1.6b 58.2  1.8b 80.0  0.2a 78.2  0.3a 0.005
Ash 1.6  0.0b 1.6  0.0b 2.7  0.0a 2.6  0.0a 0.002
Proteins 25.5  0.0a 25.1  0.1b 13.1  0.0d 13.8  0.1c <0.001
Fat 3.1  0.5a 3.4  0.7a 0.9  0.0b 1.1  0.2b 0.034
Cholesterol 170.5  5.2a 169.7  13.7a 84.6  4.3b 85.9  4.9b <0.001
Energy 192.3  1.4a 186.1  3.2a 82.1  3.2b 90.1  1.5b <0.001

Abbreviations: MY, claw muscle yield (%); EC, edible content (%), HI, hepatosomatic index; GSI, gonadosomatic index; SF, females from the Scottish coast; EF, females from the
English Channel; SM, males from the Scottish coast; EM, males from the English Channel. In each row different letters indicate significant differences between sexes and
locations (p < 0.05). The number of replicates for each location and sex were muscle (n = 10); hepatopancreas (n = 10), ovaries (n = 6) and testis (4).
720 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725

Table 4
Fatty acid profile (% of total fatty acids) of edible tissues of crabs from each location and sex (average  standard deviation).

SF EF SM EM p

Muscle
14:0 1.2  0.2a 0.53  0.10b 1.1  0.1a 0.54  0.13b 0.000
15:0 0.53  0.10a 0.56  0.07a 0.57  0.09a 0.56  0.09a 0.836
16:0 9.5  0.7a 9.0  0.3a 9.6  0.6a 10.9  0.3a 0.052
18:0 5.1  0.5a 4.9  0.2a 4.4  0.5a 4.6  0.5a 0.050
SFA 17.4  0.9a 15.7  0.5b 16.7  1.0ab 17.6  0.4a 0.001
16:1n-7 4.2  0.4b 4.9  0.2b 4.6  0.9ab 5.7  0.6a 0.001
16:1n-5 0.55  0.07ab 0.64  0.08a 0.59  0.08ab 0.49  0.07b 0.002
18:1n-9 17.0  0.8a 14.7  0.6b 16.8  1.0a 15.0  1.09b 0.000
18:1n-7 5.1  0.3b 6.2  0.3a 5.3  0.5b 6.0  0.6a 0.000
20:1n-9 2.3  0.5a 1.7  0.2b 2.0  0.4ab 1.1  0.3c 0.000
MUFA 30.8  1.7a 29.8  2.1a 30.9  1.5a 29.0  1.0a 0.056
16:3n-4 1.1  0.4a 1.2  0.3a 1.5  0.2a 1.3  0.7a 0.154
16:3n-3 2.6  0.5b 2.9  0.5b 2.8  0.4b 4.2  0.8a 0.000
16:4n-3 0.22  0.04c 0.42  0.19b 0.25  0.09c 0.71  0.06a 0.000
18:2n-6 0.99  0.07a 0.73  0.08b 1.1  0.1a 0.60  0.22b 0.000
20:2n-6 1.3  0.1a 1.1  0.1a 1.1  0.3a 0.63  0.16b 0.000
20:4n-6 7.0  0.5ab 7.4  0.6a 7.5  1.0a 6.3  0.4b 0.000
20:5n-3 21.3  2.5ab 24.1  1.5a 19.5  2.3b 21.4  1.5ab 0.001
22:5n-3 1.1  0.2a 1.1  0.2a 1.2  0.4a 1.2  0.2a 0.459
22:6n-3 10.9  1.5b 10.7  0.7b 11.5  1.3ab 12.2  0.6a 0.039
PUFA 48.4  2.7a 50.7  2.4a 48.9  1.8a 50.5  1.8a 0.095
n-3 36.6  2.8a 36.5  1.8a 35.8  2.4a 35.4  1.9a 0.648
n-6 9.6  0.5ab 10.1  1.0a 10.3  1.3a 8.8  0.6b 0.006
(n-3/n-6) 3.8  0.3a 3.6  0.5a 3.5  0.5a 4.0  0.4a 0.072
PUFA/SFA 2.8  0.2a 3.2  0.2a 2.9  0.3a 2.9  0.1a 0.053
IA 0.19  0.01a 0.15  0.01b 0.18  0.01a 0.18  0.01a 0.000
IT 0.12  0.01a 0.11  0.00a 0.12  0.01a 0.12  0.01a 0.058
Hepatopancreas
14:0 4.3  0.7a 2.7  0.8b 3.3  0.7ab 2.4  0.5b 0.000
15:0 0.72  0.11b 1.2  0.2a 0.79  0.20b 0.84  0.41ab 0.001
16:0 12.6  1.22b 12.57  1.1b 12.8  1.2ab 14.7  1.4a 0.013
18:0 3.2  0.7a 3.0  0.1a 2.7  0.6a 3.2  0.1a 0.105
SFA 21.8  2.6a 20.5  1.8a 20.5  1.8a 22.0  2.0a 0.390
16:1n-7 9.0  0.8ab 9.4  2.3ab 8.8  1.6b 11.6  1.1a 0.021
16:1n-5 0.27  0.03c 0.93  0.18a 0.25  0.07c 0.56  0.15b 0.000
18:1n-9 17.8  2.4a 17.7  3.1a 15.7  2.0ab 11.6  1.5b 0.000
18:1n-7 5.7  0.5ab 7.0  1.5a 5.3  0.7b 6.5  1.1ab 0.009
20:1n-9 6.7  1.3a 5.0  0.8a 4.6  0.8a 2.6  0.1b 0.005
MUFA 46.3  5.3a 44.7  6.9ab 38.8  4.5b 35.8  3.2b 0.003
16:3n-4 1.0  0.1a 1.0  0.1a 1.1  0.2a 1.1  0.2a 0.513
16:3n-3 0.52  0.10a 0.6  0.07a 0.56  0.12a 0.50  0.02a 0.533
16:4n-3 1.2  0.4b 1.6  0.7b 1.7  0.5b 4.2  1.2a 0.001
18:2n-6 0.91  0.07a 0.79  0.13ab 0.90  0.13a 0.62  0.10b 0.000
20:2n-6 1.6  0.2a 1.3  0.3ab 1.6  0.7a 0.67  0.15b 0.001
20:4n-6 2.6  0.5a 3.2  0.7a 3.7  1.6a 3.1  0.5a 0.193
20:5n-3 5.5  1.5b 7.0  2.6ab 7.4  1.7ab 10.3  2.0a 0.002
22:5n-3 0.8  0.2b 0.95  0.31b 1.3  0.3a 1.1  0.2ab 0.002
22:6n-3 6.5  2.1b 7.9  2.4ab 10.4  1.3a 9.3  0.7ab 0.005
PUFA 24.2  4.2b 28.0  8.2ab 32.8  2.4a 35.0  3.6a 0.002
n-3 15.4  4.2b 17.8  6.6ab 21.5  2.7ab 23.3  2.8a 0.010
n-6 5.8  0.7a 6.2  0.8a 6.9  2.3a 5.9  0.4a 0.512
(n-3/n-6) 2.6  0.7b 2.8  0.6b 3.5  1.5ab 3.9  0.3a 0.011
PUFA/SFA 1.1  0.3b 1.4  0.5ab 1.6  0.2ab 1.7  0.3a 0.001
IA 0.44  0.09a 0.34  0.07b 0.39  0.05ab 0.38  0.05ab 0.034
IT 0.27  0.06a 0.23  0.05a 0.21  0.04a 0.22  0.03a 0.061
Gonads
14:0 1.9  0.4a 1.6  0.0a 0.61  0.03b 0.86  0.42ab 0.026
15:0 0.89  0.15a 0.80  0.03ab 0.43  0.02c 0.49  0.16bc 0.002
16:0 11.7  1.1a 11.3  0.4a 8.7  0.4a 10.7  3.0a 0.051
18:0 2.5  0.1b 2.5  0.0b 5.5  0.4a 6.1  0.6a 0.036
SFA 17.6  1.8a 16.8  0.5a 16.6  0.9a 19.4  4.3a 0.054
16:1n-7 0.16  0.02a 0.14  0.01a 0.02  0.00b 0.05  0.00b <0.001
16:1n-5 0.59  0.02b 0.69  0.11a 0.63  0.02ab 0.58  0.12ab <0.001
18:1n-9 17.3  0.5a 17.0  0.7a 13.0  0.5b 11.5  1.7b 0.033
18:1n-7 4.8  0.4a 5.5  0.4a 5.2  0.2a 5.4  0.6a 0.149
20:1n-9 2.4  0.2a 2.0  0.1b 2.4  0.2a 1.7  0.0b 0.005
MUFA 34.3  0.7a 35.0  0.7a 32.2  2.1ab 30.0  0.2b 0.015
16:3n-4 0.98  0.30a 1.1  0.1a 0.44  0.01b 1.1  0.2a 0.008
16:3n-3 0.51  0.03b 0.55  0.01b 1.2  0.1a 1.0  0.2a <0.001
16:4n-3 2.4  0.3ab 1.9  0.1bc 1.5  0.1c 2.9  0.1a <0.001
18:2n-6 0.87  0.02a 0.73  0.04b 0.64  0.03bc 0.50  0.10c <0.001
20:2n-6 0.69  0.09b 0.60  0.03b 1.3  0.1a 0.54  0.01b <0.001
20:4n-6 4.9  0.2b 4.9  0.3b 9.2  0.4a 5.7  0.0b <0.001
20:5n-3 13.9  0.4b 14.8  1.0b 19.7  0.9a 18.3  0.9a <0.001
 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725 721

Table 4 (Continued )

SF EF SM EM p

22:5n-3 1.5  0.2a 1.6  0.1a 1.12  0.0b 0.85  0.10b <0.001
22:6n-3 14.2  1.6ab 13.8  0.1ab 15.5  0.9a 11.2  1.5b <0.001
PUFA 43.2  2.4a 42.6  1.3a 48.6  3.6a 43.1  0.7a 0.054
n-3 31.6  2.0b 31.7  1.2b 38.2  1.8a 32.2  2.2b 0.003
n-6 7.6  0.3b 7.6  0.3b 11.9  0.5a 8.8  1.4b <0.001
(n-3/n-6) 4.2  0.1a 4.2  0.0a 3.2  0.0b 3.7  0.3b 0.004
PUFA/SFA 2.5  0.4ab 2.5  0.2b 2.9  0.1a 2.3  0.5b <0.001
IA 0.26  0.04a 0.24  0.01ab 0.14  0.01b 0.20  0.08ab 0.048
IT 0.14  0.02a 0.13  0.01a 0.11  0.00a 0.15  0.04a 0.100

Abbreviations: SF, female crabs from the Scottish coast; EF, female crabs from the English Channel; SM, male crabs from the Scottish coast; EM, male crabs from the English
Channel; SFA, saturated fatty acids; MUFA, monounsaturated fatty acids; PUFA, polyunsaturated fatty acids; IA, index of atherogenicity; IT, index of thrombogenicity. In each
row different letters indicate significant differences between sexes and locations (p < 0.05). The number of replicates for each location and sex were muscle (n = 10);
hepatopancreas (n = 10), ovaries (n = 6) and testis (4).

aminoacids than females like histidine, lysine and arginine. The
only difference observed between populations was detected for
proline, with EC crabs showing higher concentration than SC crabs.
All identified amino acids in female gonads from both locations had
higher concentration than males on a wet weight basis. The amino
acid concentration on a wet weight basis showed marked
differences mainly between hepatopancreas and remaining
tissues, independently of sex and location (Table 7), i.e. the
concentration of total amino acids was lower in hepatopancreas
than in muscle and gonads, except for taurine and histidine. The
EAA/NEAA ratio was lower in muscle. All tissues had an excellent S-
score for all amino acids except methionine (Table 8). Hepatopan-
creas had the lowest methionine concentration, especially females,
followed by testis and muscle. Ovaries were a good source of this
amino acid.
4. Discussion
The nutritive and commercial value of seafood depends on the
chemical composition of the muscle, gonad, liver, etc., their
proportion in the specimen weight, and factors related to fishing
(Sikorski et al., 2004). However, most studies concerning the
benefits of seafood in general, and crustaceans in particular,
consider only the muscle as edible tissue (Sikorski et al., 2004).
Since the brown meat of C. pagurus (hepatopancreas and gonads) is
highly appreciated in southern Europe, it is fundamental to address
the nutritional quality of crabs considering all edible parts in order
to make wise decisions about consumption. Moreover, differences
in price that result from fishing ground location also need to be
further clarified.
As far as the meat yield is concerned, no differences were
observed between either fishing ground location; however, this
study confirmed that males, independently of location, contributed
with a higher proportion of white meat from claws than females of
the same size. This difference is due to sexual dimorphism, in
which males have bigger claws than females (Edwards, 1979); this
is used to justify the higher price of males. On the other hand,
females were responsible by a larger proportion for gonads,
although with greater variation in the gonadosomatic index due to
the different maturation stages found.

Table 5
Fatty acid profile (% of total fatty acids) of each edible tissue of crabs from both location and sexes (average  standard deviation).

Muscle Hepatopancreas Gonads p

14:0 0.86  0.34b 3.33  0.97a 1.24  0.65b <0.001

15:0 0.56  0.09b 0.88  0.28a 0.65  0.24b <0.001
16:0 9.82  0.88b 12.97  1.39a 10.45  1.72ab 0.001
18:0 4.74  0.52a 3.00  0.52b 4.11  1.70ab <0.001
16:1n-7 4.85  0.82b 9.46  1.78a 0.09  0.07c <0.001
16:1n-5 0.56  0.09a 0.48  0.31b 0.62  0.06a <0.001
18:1n-9 15.96  1.36a 16.24  3.20a 14.84  2.60a 0.264
18:1n-7 5.61  0.64ab 6.03  1.17a 5.14  0.44b <0.001
20:1n-9 1.78  0.58b 5.06  1.66a 2.21  0.29b <0.001
16:3n-4 1.26  0.43a 1.04  0.15ab 0.84  0.34b 0.005
16:3n-3 3.13  0.85a 0.54  0.09b 0.83  0.34b <0.001
16:4n-3 0.40  0.23b 1.92  1.21a 2.07  0.55a <0.001
18:2n-6 0.85  0.24a 0.83  0.15ab 0.71  0.14b 0.030
20:2n-6 1.00  0.29b 1.36  0.51a 0.87  0.36b <0.001
20:4n-6 7.03  0.81a 3.12  1.01b 6.46  2.09a <0.001
20:5n-3 21.42  2.49a 7.17  2.45c 16.71  2.77b <0.001
22:5n-3 1.14  0.26ab 1.01  0.32b 1.28  0.29a 0.021
22:6n-3 11.37  1.20b 8.31  2.66c 14.06  1.82a <0.001
SFA 16.90  1.04b 21.18  2.10a 17.46  1.96b <0.001
MUFA 30.15  1.72c 42.24  6.53a 32.96  2.18b <0.001
PUFA 49.52  2.32a 29.15  6.45b 44.88  3.61a <0.001
n-3 36.08  2.24a 18.85  5.25b 33.94  3.56a <0.001
n-6 9.70  1.04a 6.21  1.33b 9.24  2.10a <0.001
(n-3/n-6) 3.76  0.44a 3.11  1.02b 3.77  0.46a <0.001
PUFA/SFA 0.20  0.05ab 0.18  0.07b 0.25  0.07a 0.002
IA 0.17  0.02b 0.39  0.08a 0.21  0.07b <0.001
IT 0.12  0.01b 0.24  0.05a 0.13  0.02b <0.001

Abbreviations: SF, female crabs from the Scottish coast; EF, female crabs from the English Channel; SM, male crabs from the Scottish coast; EM, male crabs from the English
Channel; SFA, saturated fatty acids; MUFA, monounsaturated fatty acids; PUFA, polyunsaturated fatty acids; IA, index of atherogenicity; IT, index of thrombogenicity. In each
row different letters indicate significant differences between sexes and locations (p < 0.05). The number of replicates for each tissue were muscle (n = 40); hepatopancreas
(n = 40), gonads (n = 20).
722 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725

Table 6
Amino acid concentration (g 100 g1 of wet weight) of edible tissues of C. pagurus from each location and sex (average  standard deviation).

SF EF SM EM p

Muscle
Thr 0.99  0.03a 0.98  0.00a 1.06  0.09a 0.94  0.02a 0.099
Val 0.69  0.03b 0.70  0.01b 0.77  0.04a 0.70  0.00b 0.037
Met 0.18  0.01b 0.19  0.03b 0.26  0.02a 0.24  0.03a 0.003
Ile 0.67  0.02b 0.68  0.03ab 0.75  0.04a 0.70  0.00a 0.012
Leu 1.11  0.03b 1.13  0.04b 1.25  0.06a 1.14  0.01b 0.002
Phe 0.63  0.04ab 0.63  0.01b 0.70  0.04a 0.63  0.00b 0.037
His 0.36  0.03ab 0.35  0.02b 0.40  0.03a 0.35  0.00b 0.024
Lys 0.89  0.07a 0.96  0.04a 1.00  0.07a 0.99  0.05a 0.144
Arg 1.64  0.02b 1.67  0.07ab 1.79  0.07a 1.63  0.01b 0.004
Asp 1.49  0.02b 1.54  0.02b 1.73  0.09a 1.52  0.02b <0.001
Ser 0.67  0.02b 0.68  0.01b 0.74  0.04a 0.64  0.00c <0.001
Glu 2.50  0.01b 2.49  0.07b 2.79  0.11a 2.59  0.06ab 0.008
Gly 1.34  0.04ab 1.39  0.02a 1.24  0.08b 1.09  0.03c <0.001
Ala 0.91  0.02a 0.93  0.02a 0.98  0.06a 0.92  0.01a 0.222
Tyr 0.62  0.03b 0.63  0.01b 0.71  0.04a 0.66  0.01ab 0.003
Pro 0.73  0.10a 0.73  0.04a 0.75  0.04a 0.82  0.06a 0.212
Hyp 0.39  0.06a 0.45  0.04a 0.45  0.06a 0.44  0.02a 0.333
Tau 0.33  0.03c 0.34  0.03c 0.43  0.03b 0.50  0.01a <0.001
EAA/NEAA 0.80  0.01a 0.80  0.01a 0.81  0.01a 0.80  0.01a 0.054
TAA 16.2  0.6ab 16.5  0.5ab 17.8  0.9a 16.5  0.0b 0.032
Hepatopancreas
Thr 0.75  0.12ab 0.67  0.03b 0.75  0.02a 0.79  0.02a 0.030
Val 0.55  0.09ab 0.52  0.01b 0.60  0.05a 0.65  0.02a 0.012
Met 0.03  0.03b 0.06  0.04ab 0.12  0.04a 0.16  0.04a 0.001
Ile 0.47  0.09ab 0.44  0.01b 0.52  0.03a 0.58  0.01a 0.012
Leu 0.75  0.15ab 0.70  0.01b 0.81  0.05a 0.89  0.00a 0.020
Phe 0.46  0.10ab 0.42  0.01b 0.50  0.03a 0.54  0.01a 0.046
His 0.29  0.06b 0.27  0.00b 0.35  0.02a 0.40  0.02a 0.010
Lys 0.54  0.12c 0.54  0.01c 0.60  0.02b 0.72  0.00a 0.020
Arg 0.92  0.18c 0.88  0.03c 1.08  0.11b 1.26  0.06a 0.001
Asp 0.98  0.19abc 0.90  0.01c 1.08  0.06b 1.17  0.04a 0.046
Ser 0.47  0.09ab 0.43  0.00b 0.48  0.03a 0.50  0.01a 0.045
Glu 1.30  0.25ab 1.15  0.04b 1.39  0.09a 1.51  0.02a 0.033
Gly 0.58  0.09ab 0.54  0.02b 0.61  0.04a 0.67  0.02a 0.025
Ala 0.57  0.10ab 0.53  0.01b 0.60  0.02a 0.63  0.02a 0.029
Tyr 0.52  0.10ab 0.52  0.00b 0.58  0.02a 0.63  0.02a 0.018
Pro 0.35  0.09b 0.47  0.02a 0.39  0.02b 0.46  0.04a 0.025
Hyp 0.33  0.03a 0.32  0.01a 0.28  0.05a 0.26  0.04a 0.064
Tau 0.54  0.02a 0.31  0.02b 0.58  0.06a 0.59  0.01a 0.019
EAA/NEAA 0.84  0.02b 0.87  0.03ab 0.89  0.01b 0.93  0.01a 0.001
TAA 10.4  1.9ab 9.7  0.1b 11.3  0.7ab 12.4  0.2a 0.020
Gonads
Thr 2.03  0.03a 1.99  0.03a 0.81  0.01b 0.87  0.00b <0.001
Val 1.55  0.00a 1.56  0.03a 0.68  0.02b 0.73  0.01b <0.001
Met 0.43  0.01a 0.39  0.02a 0.09  0.00b 0.12  0.01b <0.001
Ile 1.32  0.01a 1.30  0.02a 0.49  0.01b 0.53  0.01b <0.001
Leu 2.04  0.01a 1.98  0.04a 0.75  0.01b 0.79  0.01b <0.001
Phe 1.23  0.00a 1.19  0.02a 0.51  0.01b 0.52  0.01b <0.001
His 0.72  0.01a 0.72  0.02a 0.29  0.00b 0.30  0.00b <0.001
Lys 1.13  0.04a 1.18  0.03a 0.81  0.02b 0.88  0.09b <0.001
Arg 2.13  0.02a 2.06  0.06a 0.96  0.02b 1.03  0.04b <0.001
Asp 2.37  0.02a 2.26  0.04a 1.13  0.02c 1.22  0.01b <0.001
Ser 1.64  0.01a 1.59  0.03a 0.66  0.01b 0.69  0.00b <0.001
Glu 3.60  0.07a 3.45  0.01a 1.64  0.03c 1.82  0.01b <0.001
Gly 1.30  0.01a 1.34  0.04a 0.87  0.02b 0.83  0.06b <0.001
Ala 1.28  0.01a 1.28  0.04a 0.67  0.01b 0.68  0.00b <0.001
Tyr 1.41  0.01a 1.36  0.02a 0.55  0.00b 0.57  0.00b <0.001
Pro 1.30  0.06a 1.28  0.06a 0.70  0.02b 0.74  0.03b <0.001
Hyp 0.69  0.04a 0.52  0.01a 0.34  0.01b 0.37  0.02b <0.001
Tau 0.73  0.02a 0.76  0.01a 0.49  0.01b 0.52  0.02b <0.001
EAA/NEAA 0.88  0.01a 0.77  0.00b 0.90  0.00a 0.78  0.02b 0.009
TAA 26.9  0.1a 26.2  0.5a 12.4  0.2b 13.2  0.2b <0.001

Abbreviations: SF, female crabs from the Scottish coast; EF, female crabs from the English Channel; SM, male crabs from the Scottish coast; EM, male crabs from the English
Channel; Thr, threonine; Val, valine; Met, methionine; Ile, isoleucine; Leu, leucine; Phe, phenylalanine; His, histidine; Lys, lysine; Arg, arginine; Asp, aspartic acid; Ser, serine;
Glu, glutamine; Gly, glycine; Ala, alanine; Tyr, tyrosine; Pro, proline; Hyp, hydroxyproline; Tau, taurine; EAA/NEAA, essential amino acids/non-essential amino acids; TAA,
total amino acids. In each row different letters indicate significant differences per tissue (p < 0.05). The number of replicates for each location and sex were muscle (n = 10);
hepatopancreas (n = 10), ovaries (n = 6) and testis (4).

Most differences in the chemical composition were observed
between tissues and for each tissue between sexes, whereas few
differences were observed due to fishing ground location. Previous
results obtained by Barrento et al. (2009c) for C. pagurus elemental
composition revealed this species is also an excellent source of
macro (Na, Cl and Ca) and trace elements (Fe, Cu, Zn and Se).
Barrento et al. (2009c) also found that females have higher content
of trace elements, while male crabs have more macro elements,
hepatopancreas have more elemental content than muscle and
gonads, and crabs from the SC have higher elemental content than
 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725 723

Table 7 suggested previously by Barrento et al. (2008). Other studies have

Amino acid concentration (g 100 g1 of wet weight) comparison between tissues of
also reported lack of differences in some chemical compounds in
C. pagurus from both location and sexes and with the respective p values
(average  standard deviation). respect to location, e.g. cholesterol in hepatopancreas of snow
crabs, Chionoecetes opilio (Souchet and Laplante, 2007).
Muscle Hepatopancreas Gonads p
The results obtained in this study are in agreement with the
Thr 0.99  0.06a 0.74  0.07b 1.30  0.59a <0.001 common knowledge that seafood is a source of high-quality
Val 0.71  0.04a 0.58  0.07b 1.04  0.43a <0.001 protein with a well-balanced essential amino acid composition, is
Met 0.22  0.04a 0.09  0.06b 0.23  0.16a <0.001
high in essential PUFA and shows low cholesterol levels, but in this
Ile 0.70  0.04a 0.50  0.07b 0.83  0.41a <0.001
Leu 1.16  0.06a 0.79  0.10b 1.26  0.63ab <0.001 case, also showed different magnitudes according to tissue and sex.
Phe 0.65  0.04a 0.48  0.06b 0.79  0.35a <0.001 The proximate chemical composition of edible tissues generally
His 0.36  0.03a 0.32  0.06a 0.46  0.22a 0.142 reflects their physiological functions, metabolic needs and
Lys 0.96  0.07a 0.60  0.10b 0.97  0.18a <0.001
available diet (Rosa and Nunes, 2003b; Vinagre et al., 2007).
Arg 1.68  0.08a 1.03  0.18b 1.43  0.56ab <0.001
Asp 1.57  0.11a 1.03  0.14b 1.62  0.58a <0.001
Several studies have reported similar results with decapods, like
Ser 0.68  0.04a 0.47  0.05b 1.05  0.48a <0.001 the crabs Callinectes sapidus (Çelik et al., 2004), Maja brachydactyla
Glu 2.59  0.14a 1.34  0.18b 2.43  0.92a <0.001 (Bodin et al., 2007), Carcinus mediterraneus (Cherif et al., 2008) and
Gly 1.26  0.13a 0.60  0.07b 1.04  0.24a <0.001 the penaeid shrimp Fenneropenaeus indicus (Vasquez-Boucard
Ala 0.94  0.04a 0.58  0.06b 0.92  0.31a <0.001
et al., 2004). Muscle is a structural tissue with high protein content
Tyr 0.65  0.04a 0.55  0.08b 0.89  0.41ab <0.001
Pro 0.76  0.07a 0.38  0.07b 0.94  0.29a <0.001 and poor fat. Hepatopancreas is a multifunctional organ responsi-
Hyp 0.43  0.05a 0.30  0.04b 0.45  0.14a <0.001 ble for the absorption and storage of ingested materials, such as
Tau 0.40  0.08b 0.56  0.04a 0.60  0.13a <0.001 lipids and cholesterol in decapod crustaceans. (Gibson and Barker,
EAA/NEAA 0.80  0.01b 0.88  0.04a 0.82  0.06a 0.045
1979). Gonads are responsible for the development of reproductive
TAA 16.73  0.83a 10.96  1.40b 18.24  6.98a <0.001
cells and, therefore, there is a high energy investment in this organ
Abbreviations: Thr, threonine; Val, valine; Met, methionine; Ile, isoleucine; Leu, confirmed by the high protein, fat and cholesterol levels (Wen
leucine; Phe, phenylalanine; His, histidine; Lys, lysine; Arg, arginine; Asp, aspartic
et al., 2001; Çelik et al., 2004). Both muscle and female gonads were
acid; Ser, serine; Glu, glutamine; Gly, glycine; Ala, alanine; Tyr, tyrosine; Pro,
proline; Hyp, hydroxyproline; Tau, taurine; EAA/NEAA, essential amino acids/non- better sources of proteins and total amino acids than hepatopan-
essential amino acids; TAA, total amino acids. In each row different letters indicate creas. The amino acid pattern found in all tissues of C. pagurus was
significant differences per tissue (p < 0.05). The number of replicates for each tissue characterized by higher amounts of non-essential amino acids. The
were muscle (n = 40); hepatopancreas (n = 40), gonads (n = 20).
lower amino acid content of hepatopancreas is in agreement with
results found in literature for other crustaceans, like shrimps,
EC crabs. However, as far as the meat yield and chemical lobsters and crabs (Rosa and Nunes, 2003a; Naczk et al., 2004;
composition are concerned, there is no reason for price differenti- Chen et al., 2007; Barrento et al., 2009b). In general, crustacean
ation between crabs from different locations. The main reason for muscle contains more non-essential amino acids, which are
the price distinction is most probably due to a condition factor as synthesized or modified in the body, whereas this difference is
not so evident in hepatopancreas. Ovaries were a better source of
Table 8 amino acids than testis considering the wet weight. Nevertheless
Essential amino acid scores (%) of edible tissues of C. pagurus from both locations on dry bases, differences are less significant. All tissues had S-
and sexes.
scores above 100%, except for methionine. Similar results were
SF EF SM EM obtained in the muscle of Chinese mitten crab Eriocheir sinensis
Muscle
(Chen et al., 2007). High levels of taurine were found in C. pagurus
Val 117 95 96 100 hepatopancreas. Although up to now taurine still does not have
Thr 287 227 226 228 any recommended dietary intake values, the ingestion of taurine is
Phe + Tyr 220 176 181 188 beneficial as it promotes reduced cardiovascular disease risk (Park
Met 75 63 81 83
et al., 1998; Militante and Lombardini, 2004; Elvevoll et al., 2008).
Lys 134 113 109 123
Leu 125 102 104 108 Humans only have limited ability for biosynthesis of taurine by the
Ile 149 122 123 129 liver and reabsorption by kidneys, and therefore taurine needs to
His 160 123 129 129 be acquired with the diet (Militante and Lombardini, 2004).
Hepatopancreas The higher amount of MUFA detected in hepatopancreas lipids
Val 116 123 113 119
Thr 269 270 239 243
compared to muscle is similar to other decapods (Chapelle, 1977;
Phe + Tyr 212 218 210 218 Harrison, 1990; Floreto et al., 2000; Çelik et al., 2004; Latyshev
Met 13 32 54 68 et al., 2009). The major fatty acids differences found among tissues
Lys 99 111 98 115 were registered for 16:1n-7 and 20:1n-9, which might be
Leu 105 110 102 107
associated with different metabolic needs of each tissue and with
Ile 129 136 128 138
His 158 164 170 186 biosynthesis occurring in hepatopancreas, similar to results
Gonads reported by Latyshev et al. (2009) for five crab species of the
Val 156 159 134 136 north-western Pacific Ocean, namely Paralithodes camtschaticus, P.
Thr 345 345 270 275 platypus, Chionoecetes opilio, C. angulatus and C. japonicus. In
Phe + Tyr 272 268 214 207
Met 106 98 44 53
general, the fatty acid profile of gonads was similar to muscle. The
Lys 99 106 137 142 major differences were detected in the higher proportion of 16:4n-
Leu 136 134 97 97 3 and DHA in gonads but lower values of EPA. This indicates that
Ile 172 173 126 128 16:4n-3 and DHA might be important in gonads maturation. The
His 189 191 147 145
higher EPA values in muscle might be related to its physiological
Abbreviations: SF, female crabs from the Scottish coast; EF, female crabs from the function as an important structural component of cell membranes
English Channel; SM, male crabs from the Scottish coast; EM, male crabs from the (Lilly and Bottino, 1981).
English Channel; Thr, threonine; Val, valine; Met, methionine; Ile, isoleucine; Leu,
leucine; Phe, phenylalanine; His, histidine; Lys, lysine; Tyr, tyrosine. The number of
One of the major concerns about food quality and nutrition in
replicates for each location and sex were muscle (n = 10); hepatopancreas (n = 10), developed countries is the cholesterol, fat and PUFA contents.
ovaries (n = 6) and testis (4). Cholesterol content of C. pagurus muscle (37–47 mg 100 g1) was
724 S. Barrento et al. / Journal of Food Composition and Analysis 23 (2010) 716–725
lower than other commercially important decapod species, such as
the crayfish Nephrops norvegicus (97 mg 100 g1) and shrimp
Pandalus borealis (107 mg 100 g1) (Oehlenschläger, 2006). Among
all analyzed tissues, female gonads had the highest cholesterol
concentration (169.7–170.5 mg 100 g1). Nonetheless, since the
current dietary recommendations suggest a cholesterol intake
below 200–300 mg per day and C. pagurus is an occasional food
item in the diet, all tissues pose minor risks. Seafood is considered
as a low fat food like many fish species, and almost all shellfish
contains less than 2.5% of total fat in the muscle, and only a small
number of species contains more than 15% (Nunes et al., 2003).
The present study confirms that this conclusion applies only to
muscle. The fat content of C. pagurus showed marked differences
between tissues and sexes (in gonads only). The total fat in muscle
(0.2–0.4%) and gonads of both sexes (0.9–1.1%, males; 3.1–3.4%,
females) was below 5%, a value generally considered to character-
ize a low fat food, in contrast to hepatopancreas (10.0–12.2%). The
ratio of n-3/n-6 is an excellent index to compare the relative
nutritional value of lipids, where high values generally correspond
to better quality foods. Compared to other crustacean species, like
the crabs Callinectes sapidus (muscle, 2.32; hepatopancreas, 1.57)
and Carcinus mediterraneus (muscle, 1.4), all edible tissues of C.
pagurus had higher nutritional quality in respect to this ratio (Çelik
et al., 2004; Cherif et al., 2008). The United Kingdom Department of
Health stipulated an ideal n-6/n-3 ratio of 0.25 in human diet and a
recommended value of PUFA/SFA ratio of 0.45 (HMSO, 1994). In
this regard all edible tissues had ratios in the range of the
recommended values. Although hepatopancreas had the highest AI
and TI values compared to muscle and gonads, such values were
lower than those of other food items, such as lamb (1.00), beef
(0.72), pork (0.69), rabbit (0.82) and chicken (0.50) (Rosa and
Nunes, 2003a).
5. Conclusions
Most differences in the chemical composition, fatty acid and
amino acid content in the edible tissues of female and male C.
pagurus from the Scottish coast and English Channel were related
to tissue and sex. There was not clear evidence that the fishing
ground influenced the chemical composition in a significant way.
C. pagurus muscle is rich in proteins and low in fat, while the
hepatopancreas had high fat, cholesterol and energy contents.
Though gonads were rich in protein, this tissue also had high levels
of cholesterol and energy. All edible tissues were valuable sources
of essential amino acids. The fatty acid profile in muscle and
gonads showed a pattern dominated by PUFA, while hepatopan-
creas was richer in MUFA and SFA. This pattern associated with
lower n-3 content in hepatopancreas, a lower n-3/n-6 fatty acid
ratio and higher n-6/n-3 fatty acid ratio contributed to higher
atherogenic and thrombogenic indexes. Nonetheless, compared to
other food items such as poultry, all edible tissues from these crabs
can be considered as balanced and nutritious food. However,
previous studies have highlighted that Cd content in hepatopan-
creas can be above the maximum permissible concentration and
action levels, and this is a potential concern for people who
consume large quantities of crustaceans (Barrento et al., 2009a).
Therefore, it is recommended that future studies should take into
consideration separate tissues to evaluate the nutritional quality of
crustaceans, particularly those with crabs, and to perform
consumer risk-benefit assessment analyses.
Acknowledgements
The first, second and third authors acknowledge a PhD and Post-
Doc grants, respectively, from the Portuguese Foundation for
Science and Technology (FCT) (Refs. SFRH/BD/24234/2005 and
SFRH/BPD/33090/2006). The European Commission supported this
study through the Collective Research Project, ‘‘CrustaSea:
Development of best practice, grading and transportation technol-
ogy in the crustacean fishery sector’’ (Ref. COLL-CT-2006-030421).
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