Shellfish: Nutritive Value, Health Benefits, and Consumer Safety

Abstract

Shellfish is a major component of global seafood production. Specific items include shrimp, lobsters, oysters, mussels, scallops, clams, crabs, krill, crayfish, squid, cuttlefish, snails, abalone, and others. Shellfish, in general, contain appreciable quantities of digestible proteins, essential amino acids, bioactive peptides, long-chain polyunsaturated fatty acids, astaxanthin and other carotenoids, vitamin B12 and other vitamins, minerals, including copper, zinc, inorganic phosphate, sodium, potassium, selenium, iodine, and also other nutrients, which offer a variety of health benefits to the consumer. Although shellfish are generally safe for consumption, their exposure to diverse habitats, the filter feeding nature of shellfish such as oysters, clams, and mussels, and unhealthy farming and handling practices may occasionally entail health risks because of possible presence of various hazards. These hazards include pathogenic organisms, parasites, biotoxins, industrial and environmental pollutants, heavy metals, process-related additives such as antibiotics and bisulfite, and also presence of allergy-causing compounds in their bodies. Most of the hazards can be addressed by appropriate preventive measures at various stages of harvesting, farming, processing, storage, distribution, and consumption. Furthermore, consumer safety of shellfish and other seafood items is strictly monitored by international, governmental, and local public health organizations. This article highlights the nutritional value and health benefits of shellfish items and points out the various control measures to safeguard consumer safety with respect to the products.

Full text

Shellfish: Nutritive Value, Health Benefits, and
Consumer Safety
Vazhiyil Venugopal and Kumarapanicker Gopakumar
Abstract: Shellfish is a major component of global seafood production. Specific items include shrimp, lobsters, oysters,
mussels, scallops, clams, crabs, krill, crayfish, squid, cuttlefish, snails, abalone, and others. Shellfish, in general, contain
appreciable quantities of digestible proteins, essential amino acids, bioactive peptides, long-chain polyunsaturated fatty
acids, astaxanthin and other carotenoids, vitamin B12 and other vitamins, minerals, including copper, zinc, inorganic
phosphate, sodium, potassium, selenium, iodine, and also other nutrients, which offer a variety of health benefits to the
consumer. Although shellfish are generally safe for consumption, their exposure to diverse habitats, the filter feeding nature
of shellfish such as oysters, clams, and mussels, and unhealthy farming and handling practices may occasionally entail health
risks because of possible presence of various hazards. These hazards include pathogenic organisms, parasites, biotoxins,
industrial and environmental pollutants, heavy metals, process-related additives such as antibiotics and bisulfite, and also
presence of allergy-causing compounds in their bodies. Most of the hazards can be addressed by appropriate preventive
measures at various stages of harvesting, farming, processing, storage, distribution, and consumption. Furthermore,
consumer safety of shellfish and other seafood items is strictly monitored by international, governmental, and local public
health organizations. This article highlights the nutritional value and health benefits of shellfish items and points out the
various control measures to safeguard consumer safety with respect to the products.
Keywords: consumer safety, health benefits, nutritive value, proximate composition, shellfish
Introduction
Shellfish is a major component of our global aquatic food sup-
ply. Shellfish consists broadly of 2 types of animals, crustaceans
and mollusks. Crustaceans are invertebrates with segmented bod-
ies, protected by hard shells made of chitin, and include shrimp,
lobster, crayfish, crab, and krill. Mollusks are invertebrates with soft
bodies, divided into foot and visceral section. They are subdivided
into bivalves, cephalopods, and gastropods. The commercially im-
portant bivalves are mussels, oysters, clams, and scallops, while
cephalopods include squid, cuttlefish, and octopus. The gastropod
group contains abalone, sea snail, cockle, and whelks, among oth-
ers. It is estimated that the ocean is inhabited by more than 1000
species of crustaceans, 50000 species of mollusks, besides 13000
species of finfish (Nybakken 2001). Information on the nutritive
value of shellfish is generally scattered in the literature and often
only discuss composition of general seafood items, particularly fin-
fish products. This article is intended to compile this information
and discuss recent data on the proximate compositions of differ-
ent shellfish items, and to evaluate the health benefits of individual
constituents. Shellfish may be prone to various hazards arising from
their habitats and also due to other reasons. The article also briefly
CRF3-2017-0093 Submitted 4/19/2017, Accepted 9/1/2017. Authors are with
Dept. of Food Science and Technology, Kerala Univ. of Fisheries and Ocean Sci-
ences (KUFOS), Kochi, Kerala 682506, India. Direct inquiries to author Venugopal
(E-mail: vvenugopalmenon@gmail.com,venugopalmenon@hotmail.com).
describes these hazards and measures to control these hazards. The
roles of various regulatory agencies to ameliorate these hazards in
order to ensure consumer safety are also pointed out.
Availability, Handling, and Consumption of Shellfish
Global production
According to the State of World Fisheries and Aquaculture,
published by the United Nation’s Food and Agriculture Orga-
nization (FAO), in 2014, an amount of 167.2 million metric
tons (MMT) of seafood was globally available, with landings of
shrimp, American lobsters, and cephalopods at 3.5, 0.16, and 4.3
MMT, respectively (FAO 2016). In recent times, the seafood in-
dustry is facing challenges, such as concerns about sustainability,
slow stagnation of capture fisheries, rising consumer demand, and
overall safety of the products. The landing of shrimp, one of the
major shellfish commodities, has been stable since 2012 (FAO
2016). American lobster (Homarus americanus) and Norway lobster
(Nephrops norvegicus) have accounted for more than 60% of world
lobster availability, the former reaching a record catch of 160000
tons in 2014. Cephalopods are fast-growing short-lived shellfish;
squid is the main component of the cephalopods, followed by cut-
tlefish, and octopus. The argentine short fin squid (Illex argentinus)
and the jumbo flying squid (Dosidicus gigas) are the most commer-
cially landed squid species. Since 2008, catches of cuttlefishes and
octopuses have remained relatively stable at 300000 and 350000
tons, respectively. The common octopus (Octopus vulgaris), and
the cuttlefish (Sepia spp.) remain overfished. The Pacific oyster
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Shellfish nutritive value and safety . . .
(Crassostrea gigas) is an invasive species most fecund of all oysters.
The eastern oyster (Crassostrea virginica) is moving progressively
toward overfishing (FAO 2016).
Consumer demand for shellfish and other seafood has resulted
in a significant rise in their aquaculture in fresh, brackish, and
marine waters, with a total production of 73.8 MMT in 2014, at
an estimated value of U.S.$ 160 billion. This included 16.1 MMT
of mollusks composed of 104 species valued at U.S.$ 19 billion,
and 6.9 MMT of crustacean valued at U.S.$ 36.2 billion (FAO
2016). The white leg shrimp or the Pacific white shrimp (Litope-
naeus vannamei)andthefreshwaterprawnMacrobrachium rosenbergii
cultured both in fresh water as well as brackish water are the
most farmed crustaceans. In addition, other popular shellfish such
as black tiger prawn (Penaeus monodon), scallop (Pecten yesoensis),
squids (Loligo duvauceli, Doryteuthis sibogae, and Sepioteuthis spp.),
and the cuttlefishes (Sepia pharaonis, Sepia aculeate, Sepia officinalis,
and Sepia elliptica) are farmed mainly in Asian countries (FAO
2016). Abalones (Haliotis spp.) are commercially farmed in China
(Lou and others 2013). In the past 40 y, the world abalone supply
has increased 5-fold owing primarily to increased abalone farming
practices (Suleria and others 2017). The Japanese carpet shell or
Manila clam (Ruditapes philippinarum), Yesso scallop (Patinopecten
yessoensis), blue mussel (Mytilus edulis), Asian green mussel (Perna
viridis), green-lipped mussel (P. canaliculus), the scallop (P. yessoen-
sis), crab (Scylla serrata), and the clam (Anadara granosa) are other
farmed shellfish items (Kim and Venkatesan 2015). Aquaculture
is projected to grow at almost 39% annually, with an estimated
production of about 102 MMT in 2025 (FAO 2016).
General handling and processing of shellfish
Freshly harvested shellfish are highly perishable and require care
during handling, processing, and storage; the handling and pre-
processing steps vary with the species (Su and Liu 2013). Shellfish,
upon landing, need to be immediately cleaned, washed, and sub-
jected to depuration, beheading, peeling, deveining, and other
operations, depending on the items. Molluscan shellfish such as
oysters, clams, and mussels, which filter enormous quantities of
water, accumulate microorganisms and other particulate matter in
their bodies. They are immediately decontaminated by depuration
by holding them in excess of water for a couple of days, which
reduces their microbial load (Bindu and Joseph 2005). Oysters
are initially washed with water to remove mud. The washed oys-
ters are exposed to steam in special retorts or heated by infrared
heating to open the shells, which are shucked (Martin and Hall
2006). Washed lobsters are transported in ice, their claws gener-
ally kept tied to prevent injury among the animals. Cephalopods,
being highly perishable, should not be exposed to direct sun-
light or wind, and should be carefully cleaned, and quickly iced
or frozen (Kreuzer 1984). Shrimp are prepared as whole-head on,
headless-shell-on, peeled with or without deveining, cut as fan-tail
or butterfly for freezing (Chandrasekharan 1994).
Chilling in ice is the most common postharvest processing oper-
ation. It extends shelf-life by a few days depending on the shellfish
(Venugopal 2006; Gokoglu and Yerlikaya 2015). The shelf-life of
chilled products is evaluated in terms of sensory, chemical, micro-
biological, and physical parameters (Ashie and others 1996; Huss
and others 2003). The decrease in the content of anserine (and
also other dipeptides including carnosine and glutathione) could
be a freshness index for chilled lobster (Ruiz-Capillas and Moral
2004). Most temperate shellfish like shrimp, scampi, abalone, scal-
lop, and clam have chilled shelf-life ranging from 6 to 10 d, while
their warm water counterparts remain acceptable in ice for 8 to
12 d (Ashie and others 1996; Huss and others 2003). Chilled stor-
age life of whole squid and cuttlefish are 9 and 10 d, respectively
(Vaz-Pires and others 2004; Tantasuttikul and others 2011). The
common octopus (O. vulgaris) and the adductor muscle of Pa-
cific lions-paw scallop have a chilled storage life of 8 and 12 d,
respectively (Pacheco-Aguilar and others 2008). Muscle autoly-
sis is the main reason for spoilage of cephalopods (Vaz-Pires and
others 2004). The main defect in cold-stored shrimp, lobster, and
scallop is oxidation of polyphenols resulting in a darkening of the
flesh, known as “black-spot” formation, which occurs within a
few hours after harvest (Nirmal and others 2015). In lobster, fat
oxidation may cause yellow spots; blue discoloration occurs in
crabmeat as a result of the breakdown of the copper-containing
oxygen-carrying protein hemocyanin (Huss and others 2003).
Freezing by chilled air or cryogens such as liquid nitrogen is
commercially employed for international trade of high-value shell-
fish such as shrimp, lobsters, and cephalopods (Chandrasekharan
1994; Venugopal 2006; Gokoglu and Yerlikaya 2015). Quality
losses upon prolonged frozen storage are due to changes in texture,
protein functionality, lipid oxidation, flavor changes, and drip for-
mation during thawing (Venugopal 2006). The process of individ-
ually quick-freezing (IQF) allows rapid freezing of shellfish in con-
venient, ready-to-cook quantities. Shrimp, scampi tail meat, squid
fillets (mantle), squid rings, and scallops are materials for the IQF
process. Popular IQF bivalve products include vacuum-packed
half shell oysters, clams, cockles, and scallops, which may also be
coated with bread crumbs prior to freezing (Venugopal 2006).
Apart from freezing, shellfish are subjected to thermal (blanching,
boiling, grilling, steaming, pressure cooking, frying, roasting, bak-
ing, canning), dehydration (air, vacuum, freeze-drying), breading,
high hydrostatic pressure (HHP), modified atmosphere packaging
(MAP), and other treatments (Kreuzer 1984; Venugopal 2006).
Combination-treatments can offer consumer-friendly products. A
few examples are cited. Mussel in sauce is vacuum sealed, cooked,
cooled, and frozen to give a shelf-life up to 2 y (Venugopal 2006).
A cook-chill process for peeled shrimp (Penaeus indicus) consists
of dipping in 10% brine containing 5% sodium tri-polyphosphate
followed by steaming the salted product for 10 min and packag-
ing. The product has a shelf-life of up to 25 d at 3 ±0.5 °C
(Venugopal 1993). Oysters can be preserved by a combination of
cold pasteurization and HHP treatment (Muth and others 2013)
Consumption pattern of shellfish
The demand for seafood is rapidly rising all over the world,
driven by increases in populations and their rising purchasing
power. According to a recent survey, the leading drivers of seafood
consumption are nutrition, taste, and convenience, while the
main barriers are price, availability, and concern about quality
(Christensen and others 2017). In 2014, an amount of 146.3
MMT of seafood was used as human food, giving a global per
capita seafood consumption of 20.1 kg, contributing to about
20% of total average per capita intake of animal protein. The per
capita shellfish consumption in 2013 was 4.9 kg, subdivided into
1.8 kg of crustaceans, 0.5 kg of cephalopods, and 2.6 kg of other
mollusks (FAO 2016). A recent survey reported per capita seafood
consumption of 25.8 and 35 kg in the European Union (EU) and
southern Europe, respectively (Megapesca (Portugal) 2017). Con-
sumers’ interests in shellfish products encompass fresh items, eaten
raw, or minimally processed, to variously prepared (salted, smoked,
coated, canned) and ready-to-eat items (Venugopal 2006). The
global demand for shellfish is indicated by trade figures, which
showed that, while shellfish, in quantity terms, formed 38% of
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Shellfish nutritive value and safety . . .
Table 1–Protein, fat, cholesterol, PUFA contents, and amino acid score (AAS) of shellfish.
Source A (samples cooked under moist heat) Source B (raw edible portions)
Source C (raw
edible portions)
Shellfish I II III IV V I II III IV V V
Shrimp, mixed 20.9 113 1.1 195 211 22.8 – 1.7 211 590 480
Prawn (cold)a15.4 – 0.9 143 – – – – – – –
Prawn (warm)a17.6. – 0.7 – – – – – – – –
Oyster, mixed 18.8 107 3.6 90 1480 11.4 – 3.4 79 1056 690
Oyster, eastern 14.0 107 4.9 105 1345
Squid, mixed species 15.6 108 2.0 221 549 17.9 – 1.4 233 524 490
Mussel, blue 24.0 107 4.5 56 866 23.8 – 56 1212 440
Lobster, northern 21.0 113 0.6 72 866 19.0 – 0.9 146 – 370
Crab, Dungeness 19.0 113 1.1 65 300 22.3 – 1.2 76 407 310
Crab, blue 20.2 107 1.8 100 549 – – – – – 320
Crab, Alaska king 19.4 113 1.5 53 636 – – – – –
Crab, white meat, cookeda20.5 – 0.3 66 80 – – – – – –
Scallop, mixed species, raw 17.0 107 0.8 33 300 – – – 222 220
Scallop, bay and sea 23.2 – 1.4 53 196 20.5 – – –
Clam, mixed species 25.5 107 1.9 67 396 – – – – 140
Crayfish, mixed 16.6 113 1.2 133 194 16.8 – 1.2 367 –
Cuttlefish 32.5 107 1.4 224 224 32.4 – 1.4 224 268 225
aPrawn (cold water), P. borealis, cooked; prawn (warm water), P. vannamei, raw, Department of Health, UK (2013).
Columns: I, protein; II, Amino Acid Score (AAS); III, crude fat; IV, cholesterol; and V, total PUFA. Values are given as g/100 g, w. wt. for protein and fat; and mg/100 g, w. wt. for cholesterol and PUFA.
A, SELFNutritionData; B, USDA (2012); C, Dong (2001); -, not reported.
total seafood traded in 2013; in terms of value recovered, its con-
tribution was 63.7% (FAO 2016).
Shrimp, consisting of over 300 species, is the most popular
shellfish due to its unique texture and color. Consumers gener-
ally prefer light gray or gray-colored raw shrimp, and brightly
orange-colored cooked shrimp (Parisenti and others 2011). In the
United States, per capita shrimp consumption of about 1.73 kg was
recorded in 2012; almost 90% of the shrimp consumed coming
from imports (Reed and Royales 2014). Bivalves are character-
istically tender and easily digested, which make them attractive
to the consumers. The major commercial bivalve species include
the soft-shelled as well as hard-shelled clams, blue mussel, eastern
oyster, and sea scallop. The consumption of marine mussels has in-
creased steadily over the past decades (Gr ienke and others 2014).
Japan, Korea, Argentina, Taiwan, Japan, China, and Spain con-
sume cephalopods in significant quantities (Vaz-Pires and others
2004; Kim and Venkatesan 2015). Popular cephalopods include
the common cuttlefish (S. officinalis), European squid (Loligo vul-
garis), common octopus (O. vulgaris), and the musky octopus (Ele-
done moschata) (Ozogul and others 2008; FAO 2016).The popular
crabs include Portunus spp., Charybdis spp., Chionocetes spp., the
mud crab (S. s er rata), the Dungeness crab (Metacarcinus magister),
andthebrowncrab(Cancer pagurus) (Maulvault and others 2012;
Kim and Venkatesan 2015). The shellfish species consumed in Eu-
rope are oysters, mussels, king scallops, lobsters, winkles, whelks,
cockles, clams, crab, and others (Ruiz-Capillas and Moral 2004;
Barrento and others 2009a, 2009b; Gu´
eguen and others 2011).
Marine snail (Hexaplex trunculus) is popular in northern African
countries (Zarai and others 2011). Antarctic krill (Euphausia su-
perba) is distributed around the South Pole; about 500000 tons are
harvested annually by the USSR and Japan (Gigliotti and others
2008). There is recent commercial interest in abalone because it is
a culinary delicacy, besides having therapeutic value (Suleria and
others 2017).
The consumer interest in various shellfish, as indicated above,
makes significant contribution to food security. It has been recog-
nized that food security, nutrition, and food safety are inextricably
linked (World Health Organization, WHO 2015). As Jennings
and others (2016) pointed out, aquatic food security demands that
the seafood supply should not only be sustainable to meet the
needs and preferences of people, but the products are also required
to provide them nutritional benefits while posing minimal health
risks. The proximate compositions of various shellfish and their
health benefits are discussed in the following.
Proximate Composition of Shellfish
Proximate compositions of shellfish, and also those of finfish
are provided in databases. The global FAO/INFOODS database
contains proximate compositions of raw or cooked portions of
152 crustaceans and 114 mollusks (FAO/INFOODS 2016).The
database of U.S. Dept. of Agriculture (USDA) contains about
3000 raw and processed food items including shellfish (USDA
2012). Data on proximate compositions of shellfish have also been
provided by the U.S. National Marine Fisheries Service (NMFS
1987), the Dept. of Health, UK (2013), and the Food Standards
Australia and New Zealand (FSANZ 2011). Other sources in-
clude Nettleton and Exler (1992), Venugopal and Shahidi (1996),
Gopakumar (1997), Dong (2001), Venugopal (2006), and Souci
and others (2008). The major components of shellfish are the
following.
Proteins
In general, shellfish has higher protein contents than finfish. Un-
like vertebrates, raw shellfish muscle, in addition to myosin and
other myofibrillar proteins, also contains the protein, paramyosin
up to 19% (w/w), which is rich in glutamic acid. Paramyosin,
which has a molecular weight up to 258 kDa, is found in stri-
ated and smooth muscle of invertebrates, and is involved in catch
contraction (Venugopal and Shahidi 1996). The reported aver-
age protein contents (g/100 g raw meat) of various shellfish vary:
shrimp, 17.0 to 22.1; scallop, 14.8 to 17.7; squid, 13.2 to 19.6;
crab, 15.0 to 18.4; lobster, 18.2 to 19.2; krill, 12.0 to 13.0; clam,
9.0 to 13.0; mussel, 12.6 to 13.0; cuttlefish, 16.6 to 17.3; and
oyster, 8.9 to 14.3 (NMFS 1987; Venugopal and Shahidi 1996;
Gopakumar 1997; Dong 2001; Souci and others 2008; USDA
2012). Table 1 (Source A, Column 1) provides protein contents
of shellfish items cooked under moist heat. It may be noted from
the Table 1 that samples cooked under moist heat have good pro-
tein contents. Protein contents of some raw shellfish species from
Indian waters vary from 14.0% to 21.6% (Table 2).
Protein contents of shellfish have also been reported in recent
studies. Edible portions of the Asian hard clam had 9% to 12.75%
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Shellfish nutritive value and safety . . .
Table 2–Proximate composition of some shellfish species from Indian
waters.
Shellfish Scientific name
Moisture
(%)
Protein
(%)
Crude
fat (%)
Ash
(%)
Blood clam Anadara granosa 81.4 14.5 1.8 0.8
Speckled shrimp Metapenaeus
monoceros
77.4 20.0 0.7 2.1
Backwater clam Meretrix casta 71.5 15.7 2.0 2.1
Crab Scylla serata 79.2 17.5 0.2 1.6
Cuttlefish Sepia spp. 75.8 18.1 0.2 1.4
Giant freshwater
prawn
Macrobrachium
rosenbergii
78.3 21.2 0.3 0.4
Prawns, cold
water (cooked)
Pandalus borealis 82.4 16.0 0.4 1.9
Indian white
shrimp
Penaeus indicus 77.4 20.9 0.6 1.4
Green mussel Perna viridis 76.7 12.6 2.6 2.1
Lobster, flathead Thenus orentalis 75.6 21.6 0.6 2.3
Squid Logio spp. 75.0 19.9 0.9 0.5
Antarctic krill Euphasia superba 84.7 13.0 7.0 3.0
Source: Adapted from Gopakumar (1997).
proteins. Myofibrillar proteins were the major fractions of foot and
mantle of the clam, constituting 31% to 40% (Karnjanapratum and
others 2013). The breast, claw meat, and hepatopancreas of the
blue crab have about 19% protein, w. wt. (K ¨
uc¸¨
ukg¨
ulmez and oth-
ers 2006). Generally, chemical composition of brown meat (tissue
in the body cavity comprised mainly gonads and hepatopancreas)
differed significantly from muscle (white meat in claws and legs)
of crabs (Maulvault and others 2012). The crude protein contents
of claw meat and hepatopancreas of the green crab (C. mediterra-
neus) were 17.8% to 18.2% and 13% to 14%, respectively (Cherif
and others 2008). The meat of white shrimp had higher protein
contents than that of black tiger shrimp; the former having higher
amounts of stromal proteins with greater pepsin-soluble collagen
(Sriket and others 2007). The raw meat of brown shrimp (Crangon
crangon) from the Black sea has a protein content of about 18.5%
(Turan and others 2011). Marine snail meat and hepatopancreas
are important sources of protein (Zarai and others 2011). The ed-
ible portion of common octopus (O. vulgaris) has appreciable level
of proteins (Vaz-Pires and others 2004).
Free amino acids
Free amino acids (FAAs) constitute an important fraction of
nonprotein nitrogenous compounds in shellfish muscle. Crus-
taceans have a high content of amino acids in comparison with fin-
fish. The amino acids alanine, glutamic acid and glycine are greatly
responsible for the flavor of cooked shellfish. Alanine and glycine
contribute to sweet tastes, and glutamic acid to the “umami” taste
typical of crustaceans (Huss and others 2003). Shrimp has good
contents of glycine (up to 1% of the fresh muscle), alanine, and
proline (NMFS 1987). Muscle tissues of both white and black
shrimp contain arginine, leucine, isoleucine, and proline. The
major essential amino acids (EAAs) of red and pink shrimp are
arginine, lysine, leucine, and methionine, while the non-EAAs
are glutamic acid, aspartic acid, proline, and glycine (Rosa and
Nunes 2004). Glutamic acid and glycine contents were greater
in black tiger shrimp meat, while white shrimp had good levels
of hydroxproline. Arginine, leucine, isoleucine, and proline were
predominant in both shrimps (Sriket and others 2007). The brown
shrimp has high contents of EAAs, namely, leucine, iso-leucine,
valine, and lysine, while non-EAAs were aspartic acid, glutamic
acid, glycine, and alanine (Turan and others 2011). The Norway
lobster has EAAs, namely, threonine, leucine, valine, lysine, and
arginine, each at levels of 40 mg% of raw meat. It has also the
non-EAAss glycine, alanine, and glutamic acid at 57.9, 57.2, and
31.2 mg%, respectively. In addition, it also contains the dipep-
tide, anserine at 53 mg% in the raw meat (Rosa and Nunes 2004;
Ruiz-Capillas and Moral 2004). Edible portions of Asian hard clam
contains up to 188 mg of EAAs per gram, dominated by leucine
and lysine (Karnjanapratum and others 2013). Glycine is the dom-
inant amino acid in the thick shell mussel (Mytilus coruscus), while
lysine, threonine, phenylalanine, and arginine are the important
EAAs (Li and others 2010a). Edible tissues of the marine snail are
good sources of EAAs, particularly aspartic acid (Zarai and others
2011). Green crab meat is well balanced in EAA contents, except
for tryptophan (Naczk and others 2004). The contents of tau-
rine (2-aminoethane sulfonic acid) in the muscle of mussel, oyster,
cuttlefish, and squid vary between 7.5% and 11.9% of total amino
acids (Suseela Mathew, 2016 May 5. personal communication).
Lipids
Shellfish items have low crude lipid contents, generally up to 2%
(w/w), as shown in Table 1. The lipid contents of several shellfish
from Indian waters also have very low lipid contents (Table 2).
Shellfish lipids have appreciable proportions of n-3 (omega-3)
long-chain polyunsaturated fatty acids (PUFAs), particularly eicos-
apentaenoic acid (EPA) (cis-5,8,11,14,17, C20:5), and docosahex-
aenoic acid (DHA) (cis-4,7,10,13,16,19, C22:6). Generally, the
contents of PUFA are higher than those of saturated fatty acids
(SFAs), and monounsaturated fatty acids (MUFAs) (Berge and Bar-
nathan 2005). The contents of EPA and DHA in shellfish usually
range between 300 and 500 mg%, raw muscle); their contents are
generally lower than those of oily finfish such as Atlantic mackerel,
salmon, and sardine (Dong 2001; Hossain and Takahashi 2012).
Passi and others (2002) reported that 3 species of cephalopods and
6 species of crustaceans (and also teleosts) from the Mediterranean
Sea have higher contents of n-3 PUFA (16.6% to 57.1%) than the
n-6 PUFA (4.1% to 10.6%); all of the species showing an n-3 to
n-6 ratio of more than 1. Besides, total PUFAs (21.7% to 61.5%)
were the highest, followed by SFA (16.9% to 41.3%) and MUFA
(9.1% to 42.8%). The Mediterranean giant red shrimp (Aristaeo-
morpha foliacea) has good levels of n-3 PUFA, particularly EPA and
DHA (Bono and others 2012). Korean shellfish species are rich in
EPA and DHA and are low in MUFA (Surh and others 2003).
The lipids of the shrimp spp. (Parapenaeus longirostris and Aris-
teus antennatus), and Norway lobster (N. norvegicus) contain 42%
to 48% PUFA, 26% to 35% MUFA, and 23% to 27% SFA (Rosa
and Nunes 2004). The crude fat (1%, w/w) of brown shrimp (C.
crangon L) meat consisted of 33% SFA, 22% MUFA, and 29% n-
3 PUFA. The SFA and MUFA fractions have 21% palmitic acid
and 14% oleic acid, respectively. The PUFA consisted of EPA and
DHA at 41% and 32%, respectively (Turan and others 2011). The
n-3 PUFA of white shrimp (Peneaus vannamei) and black tiger
shrimp (P. m o n o d o n ) are 42% to 44% of crude lipids. PUFAs of the
white and black shrimps have DHA-to- EPA ratios of 1.05 and
2.15, respectively (Sriket and others 2007). The farmed freshwater
prawn (M. rosenbergii) and the wild marine shrimp species (Paracoc-
cidioides brasiliensis, Penaeus schimitti, and Xiphopenaeus kroyeri)have
about 1% lipids. The principal fatty acids in the marine shrimp
were C16:0, C16:1n-7, C18:0, C18:1n-7, C18:1n-9, C20:4n-6,
C20:5n-3, and C22:6n-3. The major fatty acids in the freshwater
prawn were C16:0, C17:0, C18:0, C18:1n-7, C18:1n-9, C18:2n-
6, C20:5n-3, and C22:6n-3. The data suggested differences in
fatty acid profiles of freshwater and marine species (Bragagnolo
and Rodriguez-Amaya 2001).
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Shellfish nutritive value and safety . . .
Lipid profiles of cephalopods have been examined in detail.
The mantles of the cephalopods E. moschata and S. officinalis, and
Todarodes sagittatus have about 2% crude lipid contents; neutral
lipids are 25% to 50% of total lipids. The fatty acids are rich in
EPA and DHA (Sinanogluu and Meimaroglou 1998). Ozogul and
others (2008) reported that the common cuttlefish (S. officinalis),
European squid (L. vulgaris), common octopus (O. vulgaris), and
musky octopus (E. moschata) had PUFA, SFA, and MU FA contents
of 43.6% to 56.5%, 28.2% to 35.3%, and 4.4% to 9.5%, respectively.
The highest proportions of SFA in cephalopods were myristic acid
(C14:0, about 1% to 3%), palmitic acid (C16:0, 15.5% to 25.2%),
heptadecanoic acid (C17:0, 1.1% to 2.6%), and stearic acid (C18:0,
4.3% to 10.0%). MUFA was composed of oleic acid (cis18:1n-9,
1.8% to 4.3%) and cis-11-eicosenoic acid (C20:1, 2.1% to 4.7%).
PUFA consisted of arachidonic acid (C20:4 n−6, 1.5% to 11.7%),
EPA, 7.9% to 17.0%, and DHA, 21.0% to 39.0%. There was also
about 2% linoleic acid (C18:2 n-6) (Ozogul and others 2008).
The fatty acids of the common octopus (O. vulgaris) consisted of
58.6% of n-3 PUFA (composed of 20.1% EPA and 26.3% DHA),
25.9% SFA, and 15.4% MUFA (Vaz-Pires and others 2004). The
contents of n-3 PUFA in the squids Loligo pealei and Illex illecebrosus
were above 50% of total fatty acids (Krzynowek and others 1989).
In the cuttlefish (S. lycidas), PUFA was about 45% of total fatty
acids. DHA was the dominant n-3 PUFA. The ratio of n-3 to n-6
PUFA was 2.48 (Wen and others 2015).
The mussel (M. coruscus) has higher contents of PUFA than SFA
and MUFA; with DHA and EPA comprised 12% to 18% and
10.8% to 14.6% of total fatty acids, respectively (Li and others
2010a). Snail tissues contain significant amounts of both n-3 and
n-6 PUFA. The contents of n-3 PUFA and SFA in the marine
snail meat were about 68% and 33% of the total fatty acids, respec-
tively. Comparable values were also found in snail hepatopancreas
(Zarai and others 2011). The PUFA of Asian hard clam contained
46% to 49% of total fatty acids, with 13% to 16% DHA, and 5% to
7% EPA (Karnjanapratum and others 2013). The raw hepatopan-
creas of green crab (C. mediterraneus) has about 23% lipids; its claw
meat had about 1% lipids only. The main SFAs were palmitic and
stearic acids. Palmitic acid represented 11.5% to 12.4%, and 11%
to 11.5% of the total fatty acids in the hepatopancreas and law
meat, respectively. The contents of stearic acid were 7.8% to 8.3%
and 7.0% to 7.3% in the hepatopancreas and in the claw meat,
respectively. Oleic acid was the dominant MUFA, which repre-
sented up to 15.0% to 17.7% of the fatty acids in hepatopancreas.
Arachidonic acid (20:4n-6) formed 13.5% of total fatty acids in
the tissues (Cherif and others 2008). The total fatty acids of green
crab (Carcinus maenas) had 20.7% SFA and 40.0% PUFA; the latter
was dominated by EPA and DHA, with the ratio of EPA-to-DHA
ranging from 1.6 to 2.8 (Naczk and others 2004). The PUFA con-
tents of 2 abalone species (Haliotis spp.) accounted for over 40%
of total fatty acids. The major fatty acids identified in muscle and
viscera were C16:0, C18:0, C20:4n-6, C20:5n-3, and C22:5n-3
(Lou and others 2013). Krill oil contained 27% PUFA, 20% to
33% phospholipids, and 64% to 77% nonpolar lipids (Tou and
others 2007).
Cholesterol and other sterols
Cholesterol is the main sterol present in shellfish, while other
sterols such as stigmasterol, desmosterol, stigmasterol, C-26 sterol,
and sitosterol may also be present in low amounts (Dong 2001).
Cholesterol content is independent of fat content and is com-
parable in wild and cultivated samples (Kanazawa 2001). Table 1
(Source A, column IV) gives cholesterol contents of some shell-
fish. Raw shellfish including mollusks contained cholesterol up to
19 mg% in muscle tissue (Ozogul and others 2015). Crustaceans,
bivalves, and cephalopods may contain total sterols at 150 to 250
mg%, w. wt. (NMFS 1987; Dong 2001; Turan and others 2011).
Hard clam had 70 to 210 mg% cholesterol, w. wt. (Karnjanapratum
and others 2013). During summer, the red shrimp and pink shrimp
contained cholesterol at 60.8 and 57.8 mg%, w. wt., respectively
(Rosa and Nunes 2004). Cholesterol contents of in raw meat of
squids (L. pealei and I. illecebrosus) over a 2-y period ranged about
110 to 450 mg% (Krzynowek and others 1989). Cholesterol levels
(mg/100 g) in the raw meat of shellfish from Indian markets were:
cuttlefish, 130 to 162; squid, 188 to 198; Antarctic krill, 33.7 to
103; prawn, 118 to 169; crab, 54 to 67; lobster, 220; and oyster, 160
(Mathew and others 1999). The sterol ranged from 114 mg% in
the wild shrimp P. brasiliensis to 139 mg% in the farmed freshwater
prawn M.rosenbergii (Bragagnolo and Rodriguez-Amaya 2001).
Carbohydrates
The contents of carbohydrate including dietary fiber in shellfish
tissue are low. Carbohydrate varies from 1.3% in cooked lob-
ster meat to 2% to 3% in oyster, and the green mussel (USDA
2012; FAO/INFOODS 2016). Glycogen contents of 1.0% to
1.2% were recorded dur ing winter in red shrimp, pink shrimp,
and Norway lobster (Rosa and Nunes 2004). Pacific oyster had a
glycogen content of 6.5 ±3.0%, d. wt., during winter (Dridi and
others 2007). Asian hard clam had a maximum of 7.9% carbo-
hydrates (Karnjanapratum and others 2013) The mussels (Mytilus
spp.) contained mytilan, a noncovalently linked complex of 95%
polysaccharide and 5% protein, and another polysaccharide, a
(1-4)-d-glucan (Grienke and others 2014).
Carotenoids
Carotenoids influence color of shellfish including processed
items and hence affect their consumer acceptability. Animals, in-
cluding humans, do not synthesize carotenoids de novo and rely
upon diet as the source of these compounds. Shellfish accumu-
late carotenoids in their body tissues from carotenoid-rich marine
plants, which are used as their feeds. The carotenoids can be ei-
ther hydrocarbons or xanthophylls (the oxygenated derivatives),
and include astacene, astaxanthin, canthaxanthin, cryptoxanthin,
fucoxanthin, lutein, neoxanthin, violaxanthin, zeaxanthin, allox-
anthin, and β-carotene (Shahidi and others 1998; Venugopal 2009;
de Carvalho and Caramujo 2017). The carotenoids, βcarotene
and βcryptoxanthin, ingested by the animals may be converted
to different compounds including vitamin A. Carotenoid contents
vary depending on body parts of shellfish; they are high in their
carapace, followed by head; while their meat has minimum con-
tents. For example, the total carotenoid contents (mg/g) of raw
meat, head, and carapace of shallow water shrimp (P. m o n o d o n )are
17.4, 58.4, and 86.6, respectively. The carotenoid contents (mg/g)
of raw meat, head, and carapace of some other shellfish species are:
shallow water shrimps (Metapenaeus dobsoni), 11, 51, and 83; Para-
peneopsis stylifera, 16, 153, and 104; P. indicus, 10, 36, and 60; deep
sea shrimps, Solonocera indica, 15, 68, and 116; and Arcotheres alcocki,
21, 185, and 117, respectively (Soumya and Sachindra 2015). The
carotenoid content in raw body tissues of scallop ranged from 7
to 60 μg/g. The pigment contents varied in this order: gonad >
mantle >adductor >gill (Zheng and others 2010).
Astaxanthin is the red-orange-colored carotenoid, which re-
mains either free or bound to macromolecules such as proteins
and/or chitin, in shrimp, prawn, krill, crab, and lobster. The pig-
ment is formed from β-carotene or zeaxanthin through oxidative
C2017 Institute of Food Technologists®Vol.16,2017 rComprehensive Reviews in Food Science and Food Safety 1223

Shellfish nutritive value and safety . . .
transformation (Soumya and Sachindra 2015). Astaxanthin is the
major carotenoid present in the meat and shell of Indian shrimp
(Sachindra and others 2005a) and marine and freshwater crabs
(Sachindra and others 2005b). Other shellfish carotenoids include
canthaxanthin present in crayfish, mytiloxanthin in mussel, and
mactraxanthin and fcoxanthinol present in clams (Sachindra and
others 2005a, 2005b; Grienke and others 2014). Balachandran
(1976) reported the presence of lutein and astacene, in addition to
astaxanthin, in Indian prawn (P. stylifera). The main pigment in the
muscle tissues of the Yesso scallop, one of the important farmed
scallops in China, was identified as pectenolone (3, 3-dihydroxy-
β,β-caroten-4-one) (Li and others 2010b).
Vitamins
Shellfish species contain most of the vitamins, particularly vi-
tamin B12. Typical contents are shown in Table 3. The vitamin
contents may vary significantly in crustaceans. Oyster, blue mussel,
and clam tissues are good sources of niacin and vitamin B12. Con-
tents of vitamin B12 are generally higher in muscle tissues of crab
and lobster (FAO/INFOODS 2016). Shrimp, blue mussel, oyster,
and scallop are good sources of vitamin A (NMFS 1987; Venu-
gopal and Shahidi 1996; Souci and others 2008; USDA 2012).
Shrimp recorded vitamin D3content of about 0.06 μg/100 g
(Bogard and others 2015).
Minerals
Raw shellfish species have ash contents up to 2.0%, as shown in
Table 2. Shellfish minerals contain both macroelements, (sodium
[Na], potassium [K], calcium [Ca], phosphate [Pi],and magnesium
[Mg]), and microelements (chromium [Cr], cobalt [Co], copper
[Cu], fluorine [F], bromine [Br] iodine [I], iron [Fe], selenium
[Se], zinc [Zn], and manganese [Mn]) (Anthony and others 1983;
NMFS 1987; Dong 2001; Souci and others 2008; USDA 2012).
As shown in Table 4, most shellfish are good sources of Na, K,
Pi, Fe, Zn, Se, and Cu. Indian shrimp has Na, K, Ca, Mg, and
Pi at 107, 58, 303, 250, and 176 mg/100 g raw edible meat, re-
spectively (Gopakumar 1997). Mollusks and crustaceans contain
appreciable levels of Cu and Zn (Dong 2001). Oysters are rich
in zinc, iron, and copper (Venugopal and Shahidi 1996). Recent
studies have examined mineral contents of shellfish. Mg was the
dominant mineral in white and black shrimps, followed by Ca and
Fe (Sriket and others 2007). The blue crab has significant contents
of Ca, Mg, Pi, and Na, and their contents vary in claw, breast
meat, and hepatopancreas of the crab (K¨
uc¸¨
ukg¨
ulmez and others
2006). The edible portions of clam are rich in Na, K, Ca, Mg,
Fe, Zn, and Cu (Karnjanapratum and others 2013). The meat
and hepatopancreas of snail are rich in proteins and EAAs (Zarai
and others 2011). The contents (mg/100 g) of iron, zinc, and
calcium in 55 samples of fresh fishery products including shrimp
from Bangladesh ranged from 0.34 to 19, 0.6 to 4.7, and 8.6 to
1000, respectively. Shrimp had maximum copper and iodine con-
tents of 1200 and 120 μg/100 g, respectively (Bogard and others
2015). Fresh shellfish contain iodine at values ranging from 308 to
1300 μg/kg (FAO/WHO 2004; Oehlenschl¨
ager 2012). Shrimp
shell has more iodine and bromine than raw or cooked tissues
(Mesko and others 2016).
Factors influencing proximate composition
Habitats, season, feed, species, and life cycle. Habitats, sea-
son, feed, species, and also gametogenesis and spawning cycle can
influence proximate composition of shellfish species (Berge and
Barnathan 2005; Ozogul and others 2008; Li and others 2010a,
2011). Marine shrimp have much higher total n-3 PUFA than n-6
PUFA, while most of the freshwater shrimp demonstrated much
lower total n-3 PUFA than n-6 PUFA (Li and others 2011). Bra-
gagnolo and Rodriguez-Amaya (2001) found that the fatty acid
compositions of marine shrimp (P. brasiliensis) and the popular
farmed freshwater prawn (M. rosenbergii) were different, although
both species contained total lipids of about 1.0%. Yerlikaya and
others (2013) reported that fatty acid profiles of different shrimp
species caught from deep water and shallow water varied. The
main fatty acids were C18:1n9, C16:0, C25:6n3, C22:5n3, and
C18:0. Saturated, monounsaturated, and PUFA contents of deep
water shr imp, P. longirostris and Porcellanopagurus edwardsi, and shal-
low water shrimp, Metapenaeus monoceros, were markedly different.
The shallow-water shrimp species may contain higher levels of
PUFA than their deep water counterparts, presumably due to
higher contents of PUFA-rich phytoplankton at the surface. Har-
vesting areas can influence color and lightness value of Mediter-
ranean giant red shrimp; carotenoids in the crustacean varied
with habitats (Bono and others 2012). Mineral compositions of
mollusks depend on their living environments (FAO/INFOODS
2016). Proximate composition and mineral contents of thick shell
mussel varied seasonally except for calcium and lead (Li and others
2010a). Iodine and selenium contents tend to be largely dependent
on environmental conditions (FAO/WHO 2004). The proximate
composition of 55 fishery products, including shrimp and prawn
from capture (marine and inland) and aquaculture sources, varied
depending on their habitats and species (Bogard and others 2015).
Seasonal changes in shellfish composition have been reported.
Su and others (2006) reported significantly higher levels of total
lipids including the SFA in 2 abalone species in summer, while the
contents of total n-3 and n-6 PUFAs and total MUFA were higher
in winter and spring. The major PUFAs were EPA (34% to 43%),
and docosapentaenoic acid (DPA) (40% to 53%). The contents of
DPA were significantly higher in winter, spr ing, and summer than
in autumn. A higher n-3/n-6 PUFA ratio was found in winter
and autumn in green lip abalone (Su and others 2000). Pink and
red shrimp displayed lower values for cholesterol in summer than
winter (Rosa and Nunes 2004). Glycogen contents of shrimp and
lobster varied during winter and summer, maximum glycogen
occurring in oyster between December and February (Rosa and
Nunes 2004). Proximate composition and mineral contents of the
thick shell mussel (M. coruscus) vary with season. PUFA predomi-
nated over SFA and MUFA throughout the season. DHA (12.4%
to 18.3%) and EPA (10.8% to 14.6%) were the major PUFA (Li
and others 2010a). Higher levels of lipid and PUFA were found
in Korean bivalve shellfish in early summer, with minimal values
in late summer (Surh and others 2003). Proximate composition of
the crab (C. pagurus) was affected by gender and season. During
autumn, maximum yield of brown meat was recorded. Maximum
contents of EAAs in muscle, taurine in all the tissues, EPA in
male gonads, fat and cholesterol in the crab ovaries were recorded
during autumn (Barrento and others 2009a).
Feed composition has been recognized to influence consump-
tion and utilization of the feed, determining the sustainabil-
ity of aquaculture (Jennings and others 2016). Shrimp fed with
carotenoid-rich feed has improved color and hence better con-
sumer acceptability (Parisenti and others 2011; de Carvalho and
Caramujo 2017). The shrimp (P. m o n d o n )fedwithβ-carotene-
enriched feed resulted in accumulation of astaxanthin in its muscle,
formed by the metabolic conversion of β-carotene. Besides im-
proving the shellfish color, the dietary carotenoids reduced molt-
ing cycle, enhanced growth, and caused resistance to diseases,
1224 ComprehensiveReviewsinFoodScienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®

Shellfish nutritive value and safety . . .
Table 3–Vitamin contents of some shellfish meat.
Shellfish A B1B2B6B12 E K Folic acid Niacin
Shrimpa2 51 34 130 2.0 – – 12 –
Shrimp, mixedb––––1.2–– – –
Shrimp mixedc225 IU – 0 0 1.5 1.4 0 4 2600
Shrimpd189 IU 20 15 160 1.9 1.4 1 – 3070
Lobstera– 130 88 1000 970 1500 0.1 12 –
Lobster, northernc570 IU 0 100 100 3.1 1000 0.1 11 1100
Mussela54 160 220 76 8 750 – 16 –
Mussel, blueb––––12–– –
Mussel bluec304 300 400 100 24 – – 76 3000
Cuttlefisha3 70 30 0.4 – 1500 0.1 44 2300
Cuttlefishc675 IU – 1700 300 5.4 1500 0.2 24 2200
Cuttlefishd375 IU 0.9 150 3 – 16 1216
Oystera93 160 160 200 15 850 100 7 –
Oyster, Pacificb– – –16– – – –
Oyster, eastern, – 0.4 – 29 – – – –
mixedc180 200 200 100 35 – – – 2500
Clam, mixedb– – –49– – – –
Clam, mixedc570 IU 200 400 3.4 100 – – 29 3400
Scallop mixedc100 IU 100 100 100 1.3 – –
Scallop mixedd217 IU 70 15 73 1.5 0 – – 703
Crab, bluec7 IU 100 100 200 7.3 1800 0.1 51 3300
Crab, Alaska kingd24 IU 43 43 50 11.5 – – 44 –
Crayfishc50 IU 100 100 100 2.1 – – – –
Cray fish mixedd53 IU 70 32 08 2.1 – – 44 2300
Squid, mixed speciesd33 IU 20 400 56 1.3 1.2 – 5 2200
The values are in μg/100 g tissue. Some vitamin A values are given in international units, indicated as IU.
Sources: aRaw edible portions, Souci and others (2008).
bRaw edible portions, Dong (2001).
cSamples cooked under moist heat, SELFNutritionData.
dCooked under moist heat, USDA (2012).
-, not reported.
thereby favor ing increased shrimp production (Soumya and
Sachindra 2015). Carotenoid contents of gonad and adductor
muscle of female scallop are higher than those of male (Zheng
and others 2010). The levels of tocopherols (α,β,γ,andδ)in
cultured shellfish depend on the presence in their diets (Jennings
and others 2016). Abundant availability of PUFA-rich phytoplank-
ton in the surface waters can result in higher PUFA in lipids of
shrimp (Yerlikaya and others 2013) and oyster (Dridi and others
2007; Chakraborty and others 2016b). The increase in the con-
tents of total fatty acids during autumn correlated with availability
of food rich in chlorophyll aconcentrations (Yanar and Celix
2006). Proximate composition of European lobster (Homarus gam-
marus) and American lobster (H. americanus) differed with respect
to body parts, sex, and species. Muscle and gonads were rich in
protein, whereas hepatopancreas had high fat, cholesterol, and en-
ergy contents (Barrento and others 2009b). Gametogenic cycle
has influence on biochemical composition of oyster (Dridi and
others 2007).
Influence of processing
Chilled storage, freezing, cooking, steaming, and other treat-
ments influence proximate compositions of shellfish species
(USDA 2012; FAO/INFOODS 2016). PUFA can undergo ox-
idation during prolonged frozen storage, catalyzed by minerals
such as copper and iron, present in the shellfish (Huss and oth-
ers 2003; Venugopal 2006). Ice storage can result in an increase
in moisture together with a decrease in total nitrogen content.
Furthermore, there were changes in viscoelastic properties of the
meat, suggesting denaturation of the proteins (Binsi and others
2007). Prolonged frozen storage may lead to protein denaturation
and hence result in change of the moisture content of shrimp,
with the water-to-protein ratio increasing from 5.5 to 7.5 (Saskia
and Ruth 2014).
Heat treatments such as boiling and steaming, in general, have
little impact on proximate composition of shellfish (Kreuzer 1984;
Venugopal 2006; Su and Liu 2013). Cooking, in general, dena-
tures muscle proteins and enhances their digestibility, and drastic
heating can significantly reduce the protein quality (Venugopal
2006). Thermal stability of proteins differs depending upon the
shellfish species. Muscle proteins from black tiger shrimp, espe-
cially myosin heavy chain, had higher ther mal stability than those
of white shrimp (Sriket and others 2007). Heating in the presence
of oxygen, or sun drying, resulted in oxidation of PUFA associ-
ated with the formation of highly reactive peroxides, which react
with proteins (Huss and others 2003). Cooking caused changes
in composition, leaching of water and chitin-bound carotenoids
from crab, enhancing the color of cooked meat (Maulvault and
others 2012). Thermal processing of white shrimp (L.vannamei)
resulted in up to 52% loss of astaxanthin depending on the treat-
ment conditions. The trans-astaxanthin was reduced from 32.8 to
8.7 μg/kg during microwaving, drying, and frying, while 13-cis
astaxanthin increased from 2.4 to 5.6 μg/kg. Astaxanthin diesters
had higher thermal stability than monoesters of astaxanthin with
either EPA or DHA (Yang and others 2015). Dur ing sun drying of
cooked shrimp, up to 75% of astaxanthin was degraded along with
a 6- to 8-fold increase in cholesterol oxidation products. Storage
of the cooked product for 90 d resulted in up to 83% degradation
of astaxanthin (Hern´
andez Becerra and others 2014). Carotenoids
bound to proteins are released by the action of digestive enzymes
(Babu and others 2008). Cooking of shrimp resulted in up to 43%
loss of iodine and bromine (Mesko and others 2016). Domestic
pan-frying of squid and mussel in virgin olive oil caused up to 14%
oil absorption together with an increase in squalene and MUFA
contents (Kalogeropoulos and others 2004). Frying also caused
significant loss in cholesterol and sitosterol contents (Ozogul and
others 2015). Marination reduced the phospholipid content of soft
C2017 Institute of Food Technologists®Vol.16,2017 rComprehensive Reviews in Food Science and Food Safety 1225

Shellfish nutritive value and safety . . .
Table 4–Mineral contents of some shellfish meat.
Shellfish Na K Mg Ca Pi Fe Zn Se Cu
Shrimp mixeda146 230 67 92 224 6.1 2.2 0.1 0.9
Shrimpb– – – – – 2.4 1.1 – 0.3
Tiger shrimpc176 260 58.5 107 303 2.1 1.4 0.04 –
Shrimp, mixedd546 170 37 70 237 0.5 1.6 – –
Prawns (P. borealis), cookede– – – – – – – 0.03 0.02
Shrimp, mixedf224 182 34 39 137 3.1 1.5 0.04 0.02
Lobstera270 220 24 61 234 1.0 1.6 0.1 –
Lobster spiny, mixedb– – 51 63 – 1.2 5.7 – 0.4
Lobster, northern 486 230 43 96 185 0.3 4.1 – –
Mussela296 – 30 24 200 4.2 1.8 0.1 –
Mussel, blueb– – – – – 4.0 1.6 – 1.9
Mussel, P. viridise180 251 – 64 102 0.9 – – –
Mussel bluef369 268 37 33 285 6.7 2.7 0.09 0.1
Cuttlefish, mixeda387 273 – 27 143 8.0 0.7 0.07 –
Cuttlefish, Sepia spp.e146 206 – 70 88 7.5 – – –
Cuttlefishd744 637 60 180 580 11.0 3.5 – –
Oystera160 184 32 82 157 3.3 22.0 0.03 –
Oyster, Pacificb– – – – – 5.1 16.6 – 1.6
Scallop mixedb0.3 1.0 – 0.05
Scallop, mixedd667 304 37 6 426 0.6 1.5 – –
Scallop, mixedf74 133 15 32 95 0.6 0.8 0.01 0.1
Scallop, rawg155 203 39 29 250 1.2 4.0 0.02 0.04
Clam, mixeda– – – – – 14.0 1.4 – 0.3
Clam, M. castae81 130 – 76 106 0.9 – – –
Clam, mixedf92 628 18 92 338 28 2.7 0.06 0.07
Squid, mixedd44 246 33 32 221 0.7 1.6 – 1.9
Squid mixed rawf12 70 9 9 62 0.2 0.4 0.01 0.5
Crab, bluea– – – – – 0.7 3.5 – 0.1.
Crab, S. serratae186 378 – 68 150 1.0 10.2 0.05 1.6
Crab, Alaska kingd1072 262 63 59 280 0.8 5.9 – 7.6
Crab, Alaska kingf1435 351 – 79 375 – – – –
Crayfish, mixed, cooked moist heatd97 238 33 51 241 1.1 1.5 – –
Values are given in mg/100 g or edible portions.
Sources: aSouci and others (2008).
bDong (2001).
cDayal and others (2013).
dUSDA (2012) (cooked under moist heat).
eDepartment of Health, UK (2013).
fSELFNutritionData (samples cooked under moist heat, except squid).
gFSANZ (2011).
-, not reported.
clam with partial replacement of PUFA with MUFA (Papaioan-
nou and others 2016). Thiamin is labile to heat, ionizing radiation,
and low pH conditions. Riboflavin is reasonably stable to cooking,
but is sensitive to light. Vitamin B6tends to get destroyed with
prolonged cooking. Vitamin B12 is partially degraded and loses its
biological activity during cooking and storage (Venugopal 2006).
Nutritive Value and Health Benefits of Shellfish
It is well recognized that adequate intake of nutrients is essen-
tial for good health. Several marine ingredients have been recog-
nized to possess interesting bioactivities and hence health benefits
(Abeynayake and Mendis 2014; Grienke and others 2014; Hamed
and others 2015). This section discusses the nutritive value and
health benefits of individual components of shellfish meat. An
attempt is also made to quantify nutritive value in terms of rec-
ommended dietary intake values.
Proteins
Adequate intake of nutritious protein is crucial for health. The
nutritive value of a protein is governed by its primary structure,
amino acid composition, content of EAAs, susceptibility to enzy-
matic digestion, and extent of chemical changes due to processing
such as thermal treatment (Friedman 1996). Animal feeding ex-
periments are generally employed to determine nutritive values of
proteins. These studies include the nitrogen balance method based
on protein digestibility, determination of protein efficiency ratio
(PER) (weight gained per gram of protein consumed), net protein
utilization (ratio of amino acid converted to proteins to the ratio of
amino acids intake), and biological value (a measure of absorption
and utilization of protein by the living organism) (Friedman 1996).
The protein digestibility corrected amino acid score (PDCAAS) is
based on the amino acid content of food protein, its digestibility,
and ability to supply EAAs according to requirement (Dong 2001).
The PDCAAS of shrimp is 1, indicating its good protein quality
(Dayal and other 2013). Seafood is an excellent source of pro-
teins and contains all the EAAs. The proteins are easily digested;
most proteins show a digestibility above 90% (Oehlenschl¨
ager
2012; Hamed and others 2015). Shellfish proteins have PER val-
ues that are slightly above that of casein, the major milk protein
(Venugopal 2006). Krill concentrate has approximately 78% pro-
tein, which has a PER value equal to that of casein (Gigliotti and
others 2008). The common octopus (O. vulgaris) has a biological
value of about 84 (Kreuzer 1984). Amino acid score (AAS) of
a protein is indicative of its nutritional quality, an AAS score of
100 means high protein quality (SELFNutritionData). As shown
in Table 1, proteins of shellfish species, cooked under moist heat,
have AAS scores as high as 100. The cuttlefish (S. lycidas) has AASs
above 100 for all the amino acids, except for valine (93), while
the EAA, tryptophan, has a value of 327 (Wen and others 2015).
Shellfish proteins provide all the EAAs for maintenance and g rowth
1226 ComprehensiveReviewsinFoodScienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®

Shellfish nutritive value and safety . . .
of the human body (Friedman 1996). Compared with suggested
amino acid requirements by the FAO/WHO, the hydrolysates
of the little loligo squid (Uroteuthis chinensis) has high nutritional
value, and is a potential nutritious supplement used in various food
products (Wu and others 2015). Shellfish and other seafood are
good sources of branched chain amino acids and taurine, which
act beneficially on glucose metabolism and also blood pressure
(Elmadfa and Meyer 2017). Fermented shellfish sauces are nutri-
tional condiments, which find uses in cuisines in African and south
east Asian areas (Grienke and others 2014). In view of the high
nutritional value, shellfish proteins are able to enhance the nutri-
tive value of plant proteins, which may be deficient in 1 or more
EAAs (Kim and Venkatesan 2015; Elmadfa and Meyer 2017).
Enzymatic digestion of proteins, either in vitro or in the human
digestive system, leads to formation of several peptides besides
amino acids. The digested protein can be absorbed in the intestine
in the form of single amino acids, di- or tripeptides, and oligopep-
tides (Vijaykrishnaraj and Prabhasankar 2015). Bioactive peptides
are protein fragments, which have attracted recent attention
due to their interesting physiological functions. These include
their antimicrobial, antiviral, antitumor, antioxidative, cardiopro-
tective, immune-modulatory, analgesic, antidiabetic, antiaging,
appetite-suppressing, and neuroprotective activities (Harnedy and
FitzGerald 2012; Ngo and others 2012; Abeynayake and Mendis
2014; Cheung and others 2015). Angiotensin-I-converting
enzyme (ACE) (EC3.4.15.1) plays crucial role in the regulation of
blood pressure. The enzyme promotes conversion of angiotensin-I
to the potent vasoconstrictor angiotensin II. Inhibition of ACE is
considered a therapeutic approach in the treatment of hyperten-
sion. Several shellfish-derived peptides have been recognized to
possess ACE-inhibitory activity (Harnedy and FitzGerald 2012;
Cheung and others 2015). Clam peptides have ACE-inhibitory
and hypocholesterolemic activities, bile acid-binding capacities,
and the ability to inhibit solubility of cholesterol, indicative of
their cardioprotective role (Lin and others 2010). Peptides having
antioxidant, anticoagulant, and antihypertensive properties have
also been isolated from mussel (Grienke and others 2014). A
novel anticancer peptide from the shellfish C. gigas exhibited
cytotoxic activity, inducing death of prostate, breast, and lung
cancer cells (Cheung and others 2013). Antimicrobial activities
are described in the hemolymph of spider crab, oyster, American
lobster, shrimp, and green sea urchin (Cheung and others 2015).
Indian shrimp is a source of a crustin-like putative antimicrobial
peptide (Antony and others 2010), and also an antioxidant peptide
(Gunasekaran and others 2015). Table 5 gives shellfish-derived
peptides and their physiological functions. In addition to antiox-
idant peptides, amino acids such as phenyl alanine, histidine, and
tryptophan in fish peptides may scavenge reactive oxygen species
(ROS) such as hydroxyl radical (Dean and others 1997).
Lipids
As discussed earlier, shellfish are rich in n-3 PUFA, with a ratio
of n-3 to n-6 PUFA above 1.0 (Passi and others 2002; Sriket and
others 2007). Consumption of n-3 PUFA rich shellfish increases
the ratio of n-3 to n-6 fatty acids in the body. The n-3 PUFA are
rich in EPA and DHA. Both EPA and DHA are known to play im-
portant roles in growth, development, and maintenance of health
(Gogus and Smith 2010; Hossain and Takahashi 2012). The mode
of action of the PUFAs is attributed to their ability to give rise to a
class of pharmacologically important groups of compounds, such
as prostaglandins, prostacyclins, thromboxanes, and leukotrienes
(collectively called eicosanoids). Both n-3 and n-6 PUFA are pre-
cursors of eicosanoids. Eicosanoids derived from n-6 PUFA, such
as arachidonic acid, have pro-inflammatory functions, whereas
eicosanoids derived from n-3 PUFA have anti-inflammatory prop-
erties (Calder 2014). The n-3 PUFA-derived eicosanoids antag-
onize the formation of inflammatory prostaglandin E2, derived
from arachidonic acid and other n-6 PUFA. The n-3 PUFAs
impart their anti-inflammatory effects via reduction of nuclear
factor-κB activation. This transcription factor is a potent inducer
of pro-inflammatory cytokine production. Besides, both EPA and
DHA are also able to increase secretion of adiponectin, an anti-
inflammatory adipokine (Siriwardhana and others 2012). Through
these actions, the n-3 PUFA alter cell and tissue functions that fa-
vor disease prevention and maintenance of health (Calder 2014).
There is strong evidence to suggest protective effect of n-3
PUFA including EPA and DHA on the risk of cardiovascular
disease and stroke (Kris-Etherton and others 2002; Weichselbaum
and others 2013; Hamed and others 2015). Intake of n-3 fatty acids
has also shown to lower serum cholesterol, which is beneficial to
cardiac health (Gogus and Smith 2010). A global study comprising
16 countries, 45637 individuals, 7973 cases of coronary heart dis-
eases (CHDs), 2781 fatal CHDs, and 7157 nonfatal myocardial in-
farction events suggested that long-chain n-3 PUFA are associated
with modestly lower incidence of fatal CHD (Gobbo and others
2016). Other studies also indicate that long-chain n-3 PUFA can
have anticancer, antidepression, antiaging, and antiarthritis effects.
They can also address a number of chronic diseases, including
a spectrum of liver fat-related conditions and kidney functions
(Cardoso and others 2010; Wall and others 2010; Scorletti
and Byrne 2013; Calder 2014). DHA is crucial for devel-
opment of the brain and the central nervous system in in-
fants and to suppress neuroinflammation and oxidative stress
(Abeynayake and Mendis 2014). In view of their recog-
nized health benefits, professional bodies such as American
Heart Assn., Dept. of Health, U.K. Food Standards Agency
(FSA), among others, suggest daily total intake of EPA
and DHA varying from 250 to 1000 mg/d (Venugopal 2009;
Hellberg and others 2012; Weichselbaum and others 2013). In-
terests in these fatty acids as food supplements have attracted a
current global market valued at U.S.$3.9 billion for these com-
pounds (GOED 2017).
Carotenoids
In food items, reactive oxygen species (ROS) are formed en-
zymatically, chemically, and photochemically. Autooxidation of
unsaturated lipids leads to formation of peroxy radicals (ROO.).
Other ROS include superoxide anion (O2.−), hydroxyl radical
(HO.−), and alkoxy radical (RO.−), among others. Nonradical
derivatives are hydrogen peroxide (H2O2), ozone (O3) and singlet
oxygen (1O2). The hydroxyl radical is the most reactive ROS, fol-
lowed by singlet oxygen. Reactions of ROS with food components
destroy nutrients, change the functionalities of proteins, lipids, and
carbohydrates, and lead to the formation of undesirable volatile
compounds and carcinogens. Oxidative modifications of n-3 and
n-6 PUFAs result in the formation lipid oxidation products, such
as malonldehyde and others, which adversely react with food com-
ponents resulting loss of their functional properties (Choe and Min
2006). Oxidation of low-density lipoproteins (LDLs), induced by
oxidative stress (a situation arising when the balance between pro-
and antioxidants is disturbed), plays a key role in inflammation and
progression of atherosclerosis (Abeynayake and Mendis 2014).
Antioxidants are compounds that can inhibit or retard oxida-
tion either by scavenging the free radicals that initiate oxidation or
C2017 Institute of Food Technologists®Vol.16,2017 rComprehensive Reviews in Food Science and Food Safety 1227

Shellfish nutritive value and safety . . .
Table 5–Shellfish-derived bioactive peptides and their physiological functions.
Shellfish Sequence and other characteristics Functions/activities
Shrimp (protease) Le-Phe-Val-Pro-Ala-Phe ACE inhibitory
Shrimp (cryotin) Peptideb(molecular size, <10, 10 to 30, >30 kDa) Anticancer
Wakame (papain) Tyr-Asn-Lys-Leu –
Oyster Cys, Leu, Glu, Asp, Phe, Tyr, Ile, Gly Antimicrobial
American lobster Gln-Tyr-Gly-Asn-Leu-Leu-Ser-Leu-Leu-Asn-Gly-Tyr-Arg Antimicrobial
Jumbo Squid Gelatin peptideaAntioxidant
Blue mussel (fermentation) Glu-Ala-Asp-Ile-Asp-Gly-Asp-Gly-Gln-Val-Asn-Tyr-Glu-
Glu-Phe-Val-Ala-Met-Met-Thr-Ser-Lys
ACE inhibitory, anticoagulant, antioxidant
Scallop γ-Glutamyl-valyl-glycine Flavoring agent
Spider crab Proline-arginine-rich peptideaAntimicrobial
Snow crab (protamex) Cationic peptideaAnticancer
Oyster PeptideaAnticancer, immune-stimulant
Clam (thermolysin) PeptideaACE inhibitory
Clam (protamex) Various peptidesaHypocholesterrolemic effect
Oyster (thermolysin) PeptideaAntioxidant, antihypertensive
Oyster (subtilisin) Pro–Val-Met-Gly-Asp and Glu-His-Gly-Val peptides Antioxidant
Squid skin collagen PeptideaACE inhibitory
Krill Various peptidesaACE inhibitory
Naturally present peptides in raw
muscle
Crab, shrimp Crustin or crustin-like, Callinectin, Tachyplesin Antibacterial, antifungal, antiviral
Crayfish AstacidinaAntibacterial, antifungal, antiviral
Lobster Crustin or crustin-likeaAntibacterial, antifungal, antiviral
Mussel (Neutrase) Tyr-Pro-Pro-Ala-Lys Antioxidant
Mussel Mytilin, mytimycin, Myticin, PerninaAntibacterial, antifungal, antiviral
Shrimp Penaeidin, crustin-like peptide Antimicrobial, antifungal, antioxidant
Scallop γ-Glutamyl-valyl-glycine Flavoring agent
aSequence of peptide not reported.
Proteases used for hydrolysis are given in parenthesis.
Sources: Summarized from Lin and others (2010); Antony and others (2010); Harnedy and FitzGerald (2012); Wang and others (2013); Cheung and others (2013); Cheung and others (2015); Gunasekaran and
others (2015); and Vijaykrishnaraj and Prabhasankar (2015).
by breaking the oxidative chain reactions. Carotenoids are dietary
antioxidants (in addition to tocopherols, vitamin C, and polyphe-
nols including flavonoids). The antioxidant activities of carotenoids
have been attributed to the presence of conjugated double bonds
in their structures (Lordan and others 2011; Chuyen and Eun
2017). Carotenoids are able to quench singlet oxygen and act as
in vivo scavengers of ROS. The singlet oxygen scavenging ability
is the highest for β-carotene, followed by tocopherol, riboflavin,
vitamin D, and ascorbic acid (Choe and Min 2006). Further-
more, carotenoids, in general, possess anti-inflammatory proper-
ties, presumably due to their effects on intracellular signaling cas-
cades, thereby inhibiting production of inflammatory cytokines
(Kaulman and Bohn 2014).
Astaxanthin, the major shellfish carotenoid, has the ability to
protect body tissues from oxidative damage by UV-light. It also
has anti-inflammatory and cardioprotective properties. The an-
tioxidant activity of astaxanthin is higher than that of β-carotene,
lutein, lycopene, ɑ-tocopherol, and canthaxanthin. Other poten-
tial health benefits of astaxanthin include its anticancer, antiaging,
and immunostimulating activities. The carotenoid also possesses
antidiabetes properties, controls cataracts, and inactivates Gram-
negative bacteria also Helicobacter pylori, responsible for chronic gas-
tritis (Hussein and others 2005; Higuera-Ciapara and others 2006;
de Carvalho and Caramujo 2017). Canthaxanthin and β-carotene
protect macrophage receptors from ROS, while β-carotene pro-
tects neutrophils, a major class of white blood cells, which use
ROS to kill phagocytized bacteria (Abeynayak and Mendis 2014).
Vitamins and minerals
Shellfish vitamins serve interesting physiological roles Vitamin
A has a wide variety of functions, including specific roles in
vision embriogenesis, cellular differentiation, growth, reproduc-
tion, immune status, and taste sensations. Vitamin D deficiency
leads to impaired mineralization of bone due to an inefficient
absorption of dietary calcium and phosphorus, and is associated
with an increase in parathyroid hormone serum concentration
(Wardlaw and Smith 2009). Vitamin B12 (cobalamin) deficiency
may cause health disorders such as megaloblastic anemia and neu-
ropsychiatric disorders (Hamed and others 2015). The tocopherols
are powerful antioxidants (Alfonso and others 2016). Iodine avail-
able from shellfish is a key constituent of thyroid hormones (In-
stitute of Medicine, IOM 2007; Wardlaw and Smith 2009; James
2013).
The physiological roles of minerals present in shellfish have
been reported (Seafish 2017). Copper is a part of certain enzymes
that are required to prevent oxidative damage of cell membranes
and also regulate neurotransmitters. Iron is part of hemoglobin
present in red blood cells and is involved in oxygen transport.
Calcium can function as a pro-oxidant. Zinc helps in immune
functions, healing of wounds, development of bones, and the cell
membrane structure and functions. Zinc also may have a protec-
tive effect against atherosclerosis because of its anti-inflammatory
and antioxidant functions (Bao and others 2010, Nesheim and
Yaktine 2010). Selenium is essential for normal physiology, par-
ticularly that of the brain and endocrine tissues (Finley 2007). In
humans, 25 proteins containing the amino acids, seleno-cysteine
and/or seleno-methionine have been identified. These include the
glutathione family of enzymes having antioxidant functions, and
the thioredoxin reductase family, involved in cellular respiration
(Rayman 2000). The positive role of selenium in countering
prostate and colorectal cancers has also been indicated (Seafish
2017). Furthermore, selenium and its derivatives are able to protect
against mercury intoxication (Bjerregaard and Christensen 2012;
Khora 2014). Olmedo and others (2013) observed that most shell-
fish have beneficial Hg to Se ratios and Se-health benefit values.
Shellfish are low in calories due to their low lipid and carbohy-
drate contents (Nettleton and Exler 1992; Dong 2001). Cooked
100 g portions of blue crab, clam, oyster, scallop, or shrimp
1228 ComprehensiveReviewsinFoodScienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®

Shellfish nutritive value and safety . . .
provide 95 to 160 kcal only (Food and Drug Administration, FDA
2008). Cooked warm water prawn (Pandalus borealis) has a calorie
content of 68 kcal (295 kJ)/100 g, while the cold water prawn
(P. vannamei) has a slightly higher calorie content than P. borealis
(Department of Health, UK 2013). The values for cooked brown,
and white crab meats are 145 kcal (608 kJ), and 85 kcal (360 kJ)/
100 g, respectively (UK FSA 2017).
Determination of nutritive value of shellfish meat
components
Dietary guidelines are the key recommendations to the general
population on requirements of adequate nutrients within caloric
needs, necessary for maintenance of health, weight management,
and physical activity. The U.S. 2015 to 2020 Dietary Guidelines
provides consumers of different age and sex guidance to choose a
healthy eating pattern to prevent diet-related chronic diseases. The
Guidelines are designed to meet the Recommended Dietary Al-
lowances (RDAs), and the Adequate Intakes for Essential Nutrients
set by the IOM (Anonymous 2015). The Guidelines recommend
daily intake of the following amounts of nutrients by males aged
19 to 30, namely: protein, 56 g; total fat, 20 to 35 g (saturated
fat, <10% of total fat); macrominerals (in milligrams; Na, 2300;
Ca, 1000; K, 4700; Pi, 700; and Mg, 400); microminerals (in
milligrams except vitamin D and vitamin B12; Fe, 18; Zn, 11;
Mn, 2.3; Cu, 0.9; and Se, 0.055); vitamins (in milligrams; A, 900;
B1,1.2;B
2,1.3;B
6,1.3;B
12,2.4μg; niacin, 16; D, 600 IU; E,
15; K, 0.1, and folate 400); vitamin D, 800 IU; and calor ies 2400
to 3000. The daily nutritional goals for other age and sex groups
are also given in the Guidelines (Anonymous 2015).
The RDA values can be used to determine the nutritive value
of shellfish meat in terms of Percent Daily Value (%DV) of in-
dividual nutrients. For example, against the requirement of 56 g
for protein by a male adult, as mentioned earlier, consumption of
100 g cooked shrimp meat containing 23.5 g protein fulfils 42% of
his daily protein requirement. Therefore, “42” is the %DV for pro-
tein in shrimp. Table 6 gives the %DV for nutr ients present in meat
of various shellfish types, cooked under moist heat. It can be seen
that most shellfish have %DV values of about 50 for proteins. Dayal
and others (2013) calculated %DV for various nutrients present in
tiger shrimp (P. m o n o d o n ). They reported %DV values of 75 for
total EPA and DHA contents, and a value of 70 for the EAAs me-
thionine, tryptophan, and lysine present in the shellfish. Selenium
has a %DV as high as 110, suggesting the shrimp fully satisfied
the dietary requirement for this mineral (Dayal and others 2013).
A 100-g serving of shellfish, except squid, scallop, and crayfish
can provide appreciable amounts of the vitamin B12 necessary to
satisfy its dietary requirement, as shown in Table 6. Most shellfish
species also provide significant amounts of selenium and also cop-
per, as shown by their %DV values (Table 6). In a recent study,
potential contributions of nutrients from 55 species of shrimp,
and prawn (and also finfish), from diverse habitats to satisfy pub-
lic health requirements, were deter mined. Seven species, includ-
ing prawn could simultaneously satisfy ࣙ25% of recommended
nutrient intakes for 3 or more nutrients. Iodine from shrimp
and prawn satisfied more than 25% of its nutritional require-
ment (Bogard and others 2015). It has been recognized that the
nutritional impact of shellfish consumption may be greater than
the sum of the health benefits from individual nutrients (FAO/
WHO 2011).
The nutritive values of var ious shellfish species have been
pointed out in detail by various compositional studies. Although
these studies do not attempt to quantify the nutrient contents in
terms of %DV, they do point out nutritional values with respect to
shellfish species. A few examples are cited. The giant red shrimp,
as well as Norway lobster, are valuable sources of nutrients, in-
cluding proteins, antioxidants, among others, for the human diet
(Rosa and Nunes 2004). The muscle and gonads of female crab
(C. pagurus) has favorable n-3 to n-6 ratios, and a well-balanced
EAA composition (Barrento and others 2009a; Maulvault and oth-
ers 2012). American and European lobsters have nutritive values
compatible with nutritional foods (Barrento and others 2009b).
The mussel P. v i r i d i s has balanced ratios of essential to non-EAAs,
and also of n-3 to n-6 PUFA contents (Chakraborty and others
2016a). Furthermore, the consumption of mollusks can make an
important contribution to the daily dietary intake requirement of
Se, Cu, and Zn (Storelli and others 2010). Shellfish and other
seafood provide good measures of vitamin B12 and vitamin D
(Anonymous 2015). The presence of appreciable levels of PUFA
(including EPA and DHA), vitamins, minerals, and amino acids
qualifies the oyster (Crassostrea madrasensis) a potential “health”
food. The shellfish has also atherogenic and thrombogenicity in-
dices together with a good hypocholesterolemic to hypercholes-
terolemic ratio, pointing out its health benefits (Chakraborty and
others 2016b). The blue crab could be used as a dietary sup-
plement to balance human nutrition (K ¨
uc¸¨
ukg¨
ulmez and others
2006). Nutritional claims have also been made with respect to
common octopus (Vaz-Pires and others 2004) and Asian hard clam
(Karnjanapratum and others 2013).
Nutritive value of farm-raised shellfish
Farmed shellfish can have as high levels of nutrients as their
wild counterparts (Nettleton and Exler 1992; Bragagnolo and
Rodriguez-Amaya 2001). Cultivated thick-shell mussels represent
a source of health-benefiting long-chain n-3 PUFA, EAAs, and
minerals (Li and others 2010a). Abalones farmed in China have
abundant n-3 PUFAs, particularly EPA (Su and others 2000; Lou
and others 2013; Suleria and others 2017). The popular farmed
marine mussels belonging to Mytilus spp. and Perna spp. provide
good levels of proteins, EAAs, n-3 PUFAs, and minerals (Li and
others 2010a; Grienke and others 2014). As feeds are known to in-
fluence proximate composition, as pointed out earlier, the poten-
tial exists to enhance the nutritive value of farmed shellfish using
nutrient-enriched feeds. Microalgae-supplemented feed has been
shown to increase the PUFA contents in clams, oysters, and scal-
lops (Berge and Barnathan 2005). Feeding scallops with spirulina,
the brown green microalga, known to be rich in carotenoids and
n-3 PUFA, helps the development of the shellfish gonads, result-
ing in higher fecundity, hatchery rate, and also possibly increased
contents of n-3 PUFA in the shellfish (Zhou and others 1991).
Seafood, which includes shellfish and finfish, received particular
attention in the 2015 U.S. Dietary Guidelines, which observed ev-
idence of health benefits for the general population, as well as for
women who are pregnant or breastfeeding. For the general pop-
ulation, consumption of about 8 oz/wk consisting of a variety of
seafood, which will provide an average consumption of 250 mg/d
of EPA and DHA, is associated with reduced cardiac deaths among
individuals. Consumption of DHA-rich seafood is associated with
improved infant health outcomes (Anonymous 2015).
Shellfish-derived nutraceuticals
Nutraceuticals are defined as substances that may be considered
part of a food that provide health benefits, including the prevention
and treatment of disease(s) (Venugopal 2009). Shellfish constituents
such as carotenoids, PUFAs, bioactive peptides, among others
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Shellfish nutritive value and safety . . .
Table 6–Percent daily values (%DV) for protein, some vitamins, and minerals from shellfish.
Vitamins Minerals
Shellfish Protein A B1B2B12 Fe K Se Cu Zn Mn Pi
Shrimp, mixed species 42 – – – 25 17 57 10 10 2 17
Tiger shrimpa54 4 – – – 14 110 68 12 2 51
Shrimpb40 106
Oyster, mixed species 36 – – 11 584 67 – 102 378 1211 35 20
Oysterb26 – – – – 45 6 – – – – –
Squid, mixed species – 1 1 24 22 4 – 64 95 10 2 7
Mussel, blue 48 6 20 25 400 37 – 125 7 18 340 28
Lobster, mixed species 51 2 0 4 52 2 – 61 97 19 3 19
Lobsterb54 ––– – 29– – – ––
Crab, blue 40 0 7 3 122 – 57 32 26 9 21
Crab, Alaska king 39 1 4 3 192 4 – 57 59 51 2 28
Blue crabb52 ––– – 49– – – ––
Scallop,bay 40 27 4 22 17–4015 20 –34
Scallopb58 –– – – 412– – – – –
Clam, mixed species 51 11 10 25 1648 155 – 91 34 18 50 34
Clamb44 – – – – 30 13 – – – – –
Crayfish,mixed,farmed 35 13 4 30 4 –5234 12 2624
Cuttlefish 65 13 1 102 90 60 – 128 50 23 10 58
The values are given for 100 g shellfish cooked under moist heat.
Sources: SELFNutritionData; aDayal and others (2013); bFDA (2008).
-, not reported.
can be valuable nutraceuticals for the development of functional
foods, defined as foods with specific beneficial health effect beyond
simple nutrition (Hasler 1998; Venugopal and Lele 2014). Crus-
tacean polysaccharides, particularly chitin and its derivatives, have
pharmaceutical properties such as antioxidant, anti-inflammatory,
antiallergic, antitumor, antiobesity, antidiabetes, anticoagulant, an-
tiviral, immunomodulatory, cardioprotective, and antihepatopa-
thy activities, which offer potential applications as bioactive food
ingredients as well as nutraceuticals (Vo and others 2015). The
mussel polysaccharide mytilan possesses antibacterial, antioxidant,
and immune-modulating activities. In addition, another polysac-
charide, a (1-4)-d-glucan, present in mussel is known to exhibit
antioxidant activity and a protective effect on acute liver injury in
mice (Grienke and others 2014). A lipid extract of hard-shelled
mussel (M. coruscus) possesses strong anti-inflammatory activity
and has the potential to treat rheumatoid arthritis (Fu and oth-
ers 2015). The immune-strengthening properties of New Zealand
green lipped mussel extract are valuable to relieve osteoarthri-
tis, joint pain, and also atopic asthma (Emelyanov and others
2002). The high oxyradical scavenging capacity and total phe-
nolics suggest the nutraceutical potential of the mussel P. v i r i d i s
(Chakraborty and others 2016a). Abalone is a source of valuable
bioactives with antithrombotic, anticoagulant, anti-inflammatory,
antioxidant, and anticancer activities. Polysaccharides from certain
abalones have antithrombotic activity, comparable to that of hep-
arin (Suleria and others 2017). Snail flesh extract has the potential
to treat asthma and tuberculosis (Padmanabhan and Sujana 2008).
The glucan extract from China white jade snail (Achatina fulica)
and hot water extract of Isada krill have significant antioxidant ac-
tivities, suggesting their potentials as dietary antioxidant (Liao and
others 2014; Koomyart and others 2015). Ziconotide, a peptide
found in marine cone snail, is approved for analgesic use (Cheung
and others 2015). The extract of the snail (Bellamia bengalensis)is
also valuable for its hepatoprotective activity (Gomes and others
2011). Anti-inflammatory and antiarthritic dietary supplements
such as “Seatone” and “Lyprinol” from mussel are commercially
available (Grienke and others 2014). Taurine present in crustaceans
and mollusks has the potential to reduce risks of CHD, either alone
or in combination with n-3 PUFA (SELFNutritionData).
Table 7–Various hazard categories associated with shellfish.
Hazard type Description
Environmental Microbial pathogens, parasites, biotoxins,
heavy metals, and chemical pollutants
such as PCBs, dioxins, dioxin-like
polychlorinated biphenyls (dl PCBs), and
polychlorinated dibenzofurans
Intrinsic Allergens such as tropomyosin, myosin light
chains, troponins, paramyosin,
sarcoplasmic calcium-binding proteins,
arginine kinase, hemocyanins, cholesterol
Process related Antibiotics such as chloramphenicol,
sulfonamide, tetracycline, erythromycin,
streptomycin, and β-lactams used in
farming.
Additivessuchasallergycausing
metabisulfite used to control melanosis
Hazards Associated with Shellfish
A food is safe when it poses a minimal health hazard to con-
sumers. A hazard is defined as a biological, chemical, or physical
agent in food, or a condition of food with the potential to cause an
adverse health effect to the consumer. An estimate of the probabil-
ity and severity of the hazard is considered risk. Hazards associated
with shellfish encompass raw, fresh, minimally processed, pack-
aged, prepared, and stored items. The clinical symptoms of these
hazards are specific to the dose and the health status of the con-
sumer, ranging from mild to life-threatening and chronic adverse
reactions (Huss and others 2000, 2003). In recent times, consumers
have become well aware of seafood-borne hazards. Surveys have
shown that consumers are more concerned about chemical con-
taminants in foods, in comparison with microbial hazards, because
chemicals are known to cause long-term adverse effects (Kher and
others 2013). In the interest of consumer safety it is important to
evaluate the various hazards associated with shellfish.
Shellfish-associated hazards can be broadly grouped as environ-
mental, intrinsic, and process-related, as shown in Table 7. Table 8
categorizes hazards associated with various shellfish products, in
the order of decreasing risk. Shellfish items, which are consumed
raw without any cooking, are the most hazardous, while products
consumed soon after thorough heat processing pose minimum
1230 ComprehensiveReviewsinFoodScienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®

Shellfish nutritive value and safety . . .
Table 8–Shellfish hazard categories in order of decreasing risks.
Category Description Example
1 Shellfish consumed raw
without any cooking
Mollusks, including fresh and
frozen mussels, clam,
oysters
2 Nonheat processed raw
shellfish products often
consumed with additional
cooking
Fresh/frozen crustaceans
3 Lightly preserved products
(with <6% salt in water
phase, pH >5.0)
Salted marinated, fermented,
cold smoked shellfish.
4 Semipreserved products (salt
>6%) or pH <5.0 with
added preservatives
Salted, marinated shellfish,
fermented items, caviar
5 Mildly heat-processed
(pasteurized, cooked, hot
smoked) products
Precooked, breaded items.
6 Heat processed shellfish
products
Canned, retort-pouch
sterilized items.
Source: Adapted from Huss and others (2000).
microbial hazards. This section briefly discusses various shellfish-
borne hazards and measures to control these hazards.
Environmental hazards
Environmental hazards form a major class of hazards. These are
caused by exposure of shellfish to microbial pathogens, parasites,
biotoxins, heavy metals, pesticides, and other chemical pollutants
from their habitats.
Pathogenic microorganisms
Microbial hazards are important with respect to the safety of
shellfish species because they are prone to contamination by a vari-
ety of microorganisms. Major pathogenic organisms implicated in
seafood-borne diseases include Salmonella spp. such as Salmonella
enteritidis, Salmonella paratyphi, and Salmonella typhimurium, also
Shigella spp., enterohemorrhagic and cytoxin-producing strain of
Escherichia coli (E. coli serotype O157:H7), Campylobacter spp.,
Vibrio spp., Aeromonas spp., Plesiomonas spp., Yersinia enterocolit-
ica,Clostridium botulinum, and Listeria monocytogenes. Pollution of
coastal waters can result in contamination of bivalve shellfish with
human enteric viruses, such as hepatitis A virus, norovirus, cali-
civirus, and astrovirus (Lalitha and Thampuran 2006; Roldan and
others 2011; Leroi 2014; Jennings and others 2016). Table 9 in-
dicates bacterial pathogens involved in shellfish-bor ne illnesses,
minimal doses required for their infection and clinical symptoms.
Salmonella spp. including Salmonella typhi and S. paratyphi are
responsible for high mortality rates (Amagliani and others 2012).
Pathogens, including Salmonella spp. and Shigella spp., Vibrio vul-
nificus, Vibrio parahemolyticus, Vibrio cholera,andC.botulinum Ty p e
E, have been isolated from freshly caught crustaceans and mollusks,
most likely contaminated by unhygienic harvest waters (Anony-
mous 2003). Bivalves are more prone to contamination because
they are suspension feeders that filter phytoplankton, zooplankton,
viruses, bacteria, and inorganic matter from contaminated water
(Oliveira and others 2011). The risks may be further complicated,
since many of these pathogens remain viable in chilled products.
For example, the pathogenic V. parahemolyticus, V. vulnificus, and V.
cholera can grow at 8 °C (Leroi 2014). V. parahemolyticus, a natural
inhabitant in estuarine marine water, has been recognized as the
leading causative agent of seafood-borne illness (Wang and oth-
ers 2015). Hu and Chen (2016) observed that the resistance of
these pathogenic microorganisms to antibiotics can further add to
risk. The authors observed that 78% to 93% of strains of Vibrio
parahaemolyticus isolated from 10 species of commonly consumed
crustaceans and other shellfish in China exhibited resistance to
ampicillin, rifampin, and streptomycin. About 75% of the isolates
displayed resistance to more than 1 antibiotic, and also tolerance
to heavy metals such as copper, lead, and cadmium.
C. botulinum can be present in marine sediments. C. botulinum
types B, E, and F are frequently found in marine animals in cold
or temperate waters. Aeromonas hydrophila, an agent of foodborne
diarrheal disease, produces a wide range of cytotoxic enterotoxins
and hemolysins (Khora 2014). Hepatitis A virus spreads through
raw or undercooked shellfish and can cause liver disease (Sanchez
2015). Annually more than 1.4 million new cases of hepatitis A oc-
cur worldwide (WHO 2010). Nonindigenous pathogens such as
L. monocytogenes and Staphylococcus aureus can be present in cooked
products, as a result of abuse of processing, handling and/or stor-
age conditions (Anonymous 2003). The habit of consumption of
raw or lightly cooked bivalves increases pathogen-associated risks
(WHO 2010; Karunasagar 2014; Khora 2014; WHO 2015).
Parasites
Food-associated parasites are recognized as a threat to food safety
and human health. Flatworms, roundworms, and protozoa can in-
fest the body of marine, freshwater, as well as farm-raised shellfish.
Roundworms, called nematodes, are the most common parasites
found in marine organisms and include Ascaris spp., Tr i c h u r i s spp.,
and Trichinella spp. Other parasites are tapeworms such as Di-
phyllobothrium spp. and trematodes (Chlonorchis spp., particularly
C. sinensis,Opisthorchis spp., Heterophyes spp., Metagonimus spp.,
Nanophyetes spp., and Paragonimus spp.) which are found in fresh-
water crustaceans. Anisakids are roundwor ms found in marine
crustaceans and cephalopods. These parasites may embed in the
intestinal wall of shellfish, and can be transmitted to humans. An-
imal parasites, such as Cryptosporidium spp., may also contaminate
shellfish harvesting waters. Increasing globalization of the food
supply, the trend of consuming raw, and the general ignorance
about parasites are responsible for this hazard (Higashi 1985; Ga-
jadhar 2015).
Biotoxins
Harmful algal bloom, popularly known as “red tide,” may oc-
cur in the coastal waters, and may be quite hazardous due to
the proliferation of poisonous algae such as dinoflagellates and
diatoms. Shellfish, while feeing on these algae, accumulate their
toxins. Toxin-contaminated shellfish may be found in temperate
and tropical waters, typically after the “red tide.” The Codex Al-
imentarius Standard has classified algal toxins into 5 groups, based
on their chemical structures: (i) saxitoxin (STX), (ii) okadaic acid
(OA, a polyether toxin), (iii) domoic acid (DA, a cyclic amino
acid), (iv) brevetoxin (BTX, a cyclic polyether toxin), and (v)
azaspiracid (AZA, a polyether toxin). An additional toxin group
was also considered, tetrodotoxin (TTX), due to its emergence in
shellfish. These toxins are known to cause 4 types of syndromes:
(i) paralytic shellfish poisoning (PSP), (ii) diarrhetic shellfish poi-
soning (DSP), (iii) amnesic shellfish poisoning (ASP), and (iv)
neurotoxic shellfish poisoning (NSP). PSP is caused by STX and
TTX groups; DSP by OA and AZA groups; ASP by DA group;
and NSP by BTX group (FAO 2004). PSP is associated with
the consumption of clams, mussels, oysters, scallops, lobsters, and
crabs, while scallops, clams, and blue mussels can be carriers of
DSP. NSP can be present in oysters, clams, mussels, and cockles.
Another toxin, yessotoxin, has been found in scallop and mus-
sel (FAO 2004). Consumption of toxin-contaminated seafood by
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Shellfish nutritive value and safety . . .
Table 9–Seafood borne illnesses associated with bacterial pathogens.
Pathogenic bacteria Seafood vector Minimal dose for infectionaClinical symptoms
Vibrio paramolyticus Crustaceans 105to 106Diarrhea, nausea, vomiting
Vcholera Shellfish 102to 106Abdominal pain, vomiting
diarrhea, dehydration, and
possible death
Clostridium botulinum type E Shellfish, smoked 0.01 to 1.0 mg toxin
per gram
Paralysis, diarrhea, death
C. perfringens Sporadic incidences 105to 108Diarrhea, seldom lethal
Aeromonas hydrophila Shellfish 105to 106Vomiting, diarrhea
Listeria monotycogenes Raw seafood, smoked, salted >102Diarrhea, vomiting, nausea
Bacillus cereus Seafood, squid, prawn 106to 109Diarrhea, nausea, vomiting
Salmonella spp Shrimp, mollusks >102Fever, headache, nausea,
vomiting, abdominal pain, and
diarrhea
Shigella Mollusks 101to 102Severe diarrhea, cramps, vomiting
Yersinia enterocolitica Shellfish 107to 109Diarrhea, vomiting, fever
Escherichia coli Shellfish 101to 109, depends
on strain
Diarrhea, fever
Staphylococcus aureus Contamination from infected
persons
105to 106Diarrhea, cramps, vomiting
aColony forming units/g raw shellfish meat.
Source: Adapted from Lalitha and Thampuran.(2006).
humans primarily results in acute gastrointestinal and neurolog-
ical manifestations and leading to allergic reactions. These have
been attributed to the effects of the toxins on sodium and calcium
channels, the enzymes, sodium–potassium ATPase, and protein
phosphatises (Garthwaite 2000; Kalidas and Anand 2006; Wang
2008; Khora 2014). DSP toxins in shellfish may be capable to in-
crease cancer risk (Hernandez and others 2008). Annually, more
than 50000 algal toxin-related incidents with an overall mortality
rate of 1.5% have been reported globally (Wang 2008). The lethal
effect of toxins is expressed as mouse unit, which is defined as the
minimum amount of purified toxin (in micrograms) required to
kill mouse of 20 g in 1 min, when 1.0 mL of a solution of extract
at pH 4.0 is injected interperitoneally (Kalidas and Anand 2006).
Heavy metals
Industrial spills and sewage discards in coastal waters contam-
inate shellfish with toxic heavy metals, such as mercury (Hg),
arsenic (As), cadmium (Cd), and lead (Pb). It is well known that
chronic exposure to Hg, As, Cd, and Pb can cause adverse health
effects (WHO 2015). Mercury is ubiquitous in the environment,
which enters the air during fossil-fuel combustion, mining, smelt-
ing, solid-waste incineration, and other industrial activities. The
metal exists in biological systems in elemental (metallic), inor-
ganic (such as mercuric chloride), and organic forms (such as
methylmercury [MeHg]; and ethylmercury [EtHg]). Inorganic
mercury is converted to MeHg by microorganisms, which then
enters the food chain and is bioconcentrated in the liver, gills,
ink or skin of the seafood. Exposure to even small amounts of
MeHg by pregnant women may affect fetal neurological devel-
opment (Karunasagar 2014; WHO 2015; Seafish 2017). A recent
collaborative study involving 8 countries showed that monomethyl
mercury (MMG) in mussel, squid, and crab claw, and also some
teleosts ranged from 0.035 to 3.58 μg/g, d. wt. Mussel had lowest
MMG concentration (Valdersnes and others 2016). Global surveys
indicated that intake of MeHg from shellfish and other seafood by
women and infants in certain regions exceeded the reference value
(Sheehan and others 2014).
Arsenic exists as highly toxic organic compounds in algae, which
are consumed by shellfish. The metal interferes with several en-
zymes and causes oxidative stress as well as immune, endocrine,
and epigenetic effects (Khora 2014). Lead occurs primarily in
inorganic form in the environment. Human exposure is mainly
via food and water. Lead can cause developmental neurotoxicity in
children and cardiovascular effects in adults (European Food Safety
Authority, EFSA 2010; Khora 2014). Cadmium can cause kidney
damage, whereas lead causes neurotoxicity especially in children.
A survey showed that flesh of mollusks had Hg, Cd, and Pb at
levels of 0.44, 0.49, and 0.1 μg/g, w. wt., respectively. The cor-
responding values for the metals in raw cephalopods were 0.27,
0.50, and 0.12 μg/g. Squid meat accumulated selenium up to 1.2
μg/g raw meat; mollusks carried copper at 37.4 μg/g and zinc at
42 μg/g. Cuttlefish can accumulate significant amounts of potas-
sium, calcium, manganese, iron, copper, and zinc from polluted
marine environments (Akpan and others 2009). Octopus, squid,
and cuttlefish from the Mediterranean Sea harbored heavy metals,
their hepatopancreas showing higher concentrations than flesh.
Highest concentrations of metals were found in octopus, namely:
Hg, 0.44; Cd, 0.49; Pb, 0.10; Cu, 37.4; and Zn, 42.4 μg/g, w.
wt. The contents (μg/g, w. wt.) for cuttlefish were: Hg, 0.27;
Cd, 0.50; and Pb, 0.12, while squid tended to accumulate lower
amounts of metals, especially Hg. The contents (μg/g, w. wt)
were: Hg, 0.11; Cd, 0.30; Pb, 0.05; and Se, 1.18. Cr was uni-
formly distributed among the various species at 0.38% to 0.43
μg/g, w. wt. (Storelli and others 2010). Another study reported
that cuttlefish meat had As, Cd, and Pb at concentrations of 2.2,
0.28, and 0.02 μg/kg, w. wt., respectively, while mercury was
below the detection limit (Wen and others 2015).
Chemical pollutants
Polychlorinated biphenyls (PCBs), carcinogenic dioxins (poly-
chlorinated dibenzo-p-dioxins, PCDDs), dioxin-like PCBs (dl
PCBs), and polychlorinated dibenzofurans are by-products in the
manufacture of herbicides and pesticides. They are also formed
during volcanic eruptions, forest fires, and from incomplete com-
bustion during waste incineration. These compounds are col-
lectively referred as persistent organic pollutants (WHO 2015).
Short-term exposure of humans to high levels of dioxins may
result in skin lesions, such as patchy darkening of the skin, and al-
tered liver function. Long-term exposure is linked to impairment
of the immune system, the developing nervous system, and the
endocrine system as well as reproductive functions. Chronic ex-
posure of animals to dioxins has resulted in several types of cancer
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Shellfish nutritive value and safety . . .
(James 2013; WHO 2015). The toxicity of the PCBs and dioxin
congeners is expressed as Toxin Equivalency (TEQ) (FAO/WHO
2016). Domingo (2016) observed that cooking procedures that
release fat from a seafood product can reduce the concentrations
of these fat-soluble organic contaminants
Hazards due to intrinsic factors
Shellfish allergy. Food allergy is due to the ability of molecules
(antigens) present in the food to interact with immunological
specific antibodies in the human body. Allergy symptoms in pre-
disposed individuals depend on properties of the allergens such
as their molecular size, chemical complexity, and genetic capacity
of the host. Of the different types of allergies, Type I allergy is
mediated by food proteins (Sathe and others 2016). Shrimp, lob-
ster, and crab are common allergenic seafood items. The proteins,
namely, tropomyosin, the enzyme arginine kinase, hemocyanins,
paramyosin, myosin light chains, troponins, tropomyosin, and sar-
coplasmic calcium-binding proteins are responsible for shellfish
allergy (Fernandes and others 2015).
Content of cholesterol. Cholesterol in the body is transported
by LDL and high-density lipoprotein (HDL), present in the blood.
The presence of the LDL form of cholesterol in blood is associated
with atherosclerosis, while the HDL form is inversely related to
the development of atherosclerosis. As mentioned earlier, shellfish
generally contains significant levels of cholesterol (Table 1). There
is a general concern that consumption of shellfish may lead to
deposits of LDL in the arteries leading to atherosclerosis (Wardlaw
and Smith 2009). Cooking decreased cholesterol content of crab
meat, but did not reduce it in shrimp or oyster (Krishnamoorthy
and others 1979).
Process-related hazards
Processed shellfish products may car ry pathogenic organisms
if the processing treatment is not adequate to eliminate them.
Most cases of botulism are associated with products subjected to
inadequate thermal treatment (Leroi 2014). Besides botulism, lis-
teriosis, cholera, and hepatitis A virus may be present in inade-
quately cooked, smoked, fermented, and pickled products, which
are all usually consumed without further processing (Jay and oth-
ers 2000). Listeria outbreaks are generally attributed to chilled
ready-to-eat foods. These foods will not be generally subjected
to a thorough heat treatment and therefore allow growth of L.
monocytogenes. The majority of Listeria spp. isolated from aquatic
products belongs to serotype 1/2a (Jami and others 2014).
Melanosis (black spot formation) is a problem in shrimp, lob-
ster, and scallop that adversely affects their appearance and hence
the commercial value. Melanosis is caused by oxidation of phe-
nols by the enzyme polyphenol oxidase to quinone that eventually
polymerizes to black, high-molecular-weight melanins. Chilling
shellfish does not prevent melanosis, but slows it down only. In the
Norway lobster (N. norvegicus), black spot appears within the 1st 4 d
after capture and increases gradually throughout storage (Edmonds
2006). The common practice to control melanosis is dipping the
shellfish in dilute aqueous solution of sodium metabisulfite; how-
ever, high concentrations of this chemical are frequently used by
the processors, thereby leaving appreciable residual sulfite (SO2)in
the edible portions. The chemical has been recognized to cause al-
lergic reactions, particularly in asthma patients (Nirmal and others
2015).
Hazards related to farmed shellfish. Shellfish is generally cul-
tured in protected coastal areas that are usually under pressure from
other human activities, leading to exposure of the shellfish to var-
ious hazards (Smaal and Wijsman 2010). These hazards include
contamination of shellfish by pathogenic microorganisms, para-
sites, toxin-producing organisms, and also chemical pollutants. A
bacterial disease, known as early mortality syndrome, has been
a major cause for losses in shrimp aquaculture in Southeast Asia
caused by a unique strain of the V. parahaemolyticus (Reed and
Royales 2014). The viral disease “white spot syndrome” has con-
tinued to cause annual losses up to U.S.$ 1 billion, since its emer-
gence in the 1990s (Jennings and others 2016). Tropical coun-
tries suffer greater losses in aquaculture during disease outbreaks,
thereby presenting a major problem for food production and se-
curity (Leung and Bates 2013).
Contaminants in farmed shellfish include a range of chemicals
such as veterinary pharmaceuticals (antibiotics, parasitical treat-
ments, anesthetics), disinfectants (used to decontaminate equip-
ment and eggs), and other biocidal chemicals such as formalin
(used to control diseases), feed additives (such as flesh pigments),
among others. Use of antibiotics (including chloramphenicol, sul-
fonamide, tetracycline, erythromycin, streptomycin, β-lactams,
and sulfonamides) to prevent diseases in farms constitutes seri-
ous public health hazards leading to possible emergence of an-
tibiotic resistance (Holmstr¨
om and others 2003; Hu and Chen
2016). A recent survey showed that from 2006 to 2011 about
200 of 730 samples of retail aquacultured fishery products (in-
cluding clam and shrimp) were positive for Salmonella spp. Thirty-
eight serovars were identified in the 217 Salmonella isolates. Of the
Salmonella spp., about 70% were resistant to at least 1 antimicro-
bial drug, while 43% were multidrug resistant. Resistance of the
isolates against other antibiotics were: sulfonamides (57%), tetra-
cycline (34%), streptomycin (29%), ampicillin (24%), and nalidixic
acid (21%) (Zhang and others 2015). The Hong Kong oyster (C.
hongkongensis) is widely farmed in estuarine waters, and has been
found to accumulate Cu and Zn in their soft tissues (Gao and
Wang 2014). It is well recognized that chemical pollutants are
grossly accumulated in farm sediments, causing high levels of ac-
cumulation in bottom living animals such as shrimp (Swapna and
others 2012).
Measures to Control Consumer Hazards
Safe food is of utmost importance in the interest of public health.
Food safety can be ensured to a great extent with a combination
of technology, and stringent quality standards, coupled with con-
stant vigilance by regulatory bodies. The key elements of shellfish
biosecurity include adequate diagnostic and detection methods to
monitor pathogens, disinfection and pathogen eradication meth-
ods, applications of the Hazard Analysis Critical Control Point
(HACCP) system, Good Management Practices (GMP), practical
guidance, and appropriate legislative controls (Soares and others
2016; UK FSA 2017). HACCP is a 7-step management system that
addresses food safety through the analysis and control of biologi-
cal, chemical, and physical hazards from raw material production,
procurement, handling to manufacturing, distribution, and con-
sumption of the processed product (National Advisory Committee
on Microbiological Criteria for Foods, NACMCF 1998; Huss and
others 2000; Tzouros and Aravanitoyannis 2007).
Handling procedures for shellfish have been briefly mentioned
earlier. Depuration for 2 d significantly reduces contaminated
metals such as lead in bivalves. The health status of bivalves during
depuration can be monitored by their glycogen content (Anacleto
and others (2015). Depuration also significantly reduces contam-
inated pathogens from shellfish, although it is less effective to
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Shellfish nutritive value and safety . . .
reduce hepatitis A to a significant level (Sanchez 2015). Commer-
cial processing methods are generally inadequate to completely in-
activate hepatitis A virus. For example, cooking of clam and oyster
below 70 °C for approximately 47 s, which is employed for open-
ing up their shells, could not annihilate hepatitis A virus (Sanchez
2015). Physical, chemical, and biological intervention strategies to
control Salmonella spp., Shigella spp., Vibrio spp., L. monocytogenes,
C. botulinum, viruses such as hepatitis A, and other microorganisms
are available. These include time/temperature control, control of
pH, control of water activity, and the use of preservatives, fermen-
tation, drying, and salting (Anonymous 2003; Wang and others
2015). An integrated process of cooking and vacuum-cooling
enhances the microbiological quality of mussel (Cavalheiro and
others 2013). Growth of C. botulinum type E can be controlled by
maintaining the temperature below 3.3 °C (Jay and others 2000).
HHP treatment at 500 MPa for 2 min at 0 °C significantly reduced
nonspore-forming pathogenic microorganisms from oysters (Ling-
ham and others 2016). Exposure to ionizing radiations (gamma
rays, machine-generated electron beams, or X-rays) provides an
effective safeguard against most microbial and parasitic hazards.
Gram-negative pathogens such as Vibrio and Salmonella are com-
paratively more sensitive to radiation than Gram-positive bacteria.
These pathogens have D10 values (radiation dose required for 90%
reduction) as low as 1 kGy in comparison with high D10 values
of Gram-positive organisms. Bacterial spores such as those of C.
botulinum are more resistant to radiation than their vegetative cells.
Microorganisms in frozen samples will have higher D10 values
(Venugopal and others 1999; Sommers and Rajkowski 2011). The
U.S. FDA has amended its current food additive regulations to al-
low the use of ionizing radiation at a maximum permitted dose of
6.0 kGy to inactivate foodborne pathogens in crustaceans and
extend their shelf-life (Center for Food Safety and Applied Nu-
trition, CFSAN 2014). Irradiation at this dose (6.0 kGy) caused a
collateral reduction of hepatitis A virus levels along with Vibrio spp.
(Praveen and others 2013). Radiation treatment can also inactivate
parasites in shellfish (Sommers and Rajkowski 2011). Irradiation
can also be combined with traditional processes such as depura-
tion, thermal treatment, and others (Venugopal 2006). Candling,
trimming belly flaps, and physically removing cysts can reduce
parasitic hazards. Heating at 55 °C for 1 min or by frozen storage
for 24 h at –20 °C can inactivate parasites (NACMCF 1998;
Codex Alimentarius 2016). The hazard of sulfite allergy associated
with sulfite-treated shellfish can be addressed by using sulfite al-
ternatives such as Xyrex—Prawnfresh, Everfresh, and Melacide SC20
(Edmonds 2006).
Addressing the problem of harmful algal bloom causing presence
of toxins in shellfish requires integrated coastal zone management,
particularly regular monitoring of algae in shellfish-growing wa-
ters and closing of shell harvesting areas, whenever toxin levels in
shellfish exceed limits (Khora 2014). Marine toxins being mostly
nonprotein in nature processing operations such as cooking, smok-
ing, drying, and salting have limited scope to destroy them in the
harvested shellfish. The kinetics of thermal destruction of scallop
toxin is qualitatively similar to that of microorganisms (Indrasena
and Gill 1999). Avoiding the offending food is the best defense
for sensitive individuals to prevent allergy (Sathe and others 2016).
Heat treatment enhanced antibody reactivity to prawn allergens
including tropomyosin, myosin light chain, sarcoplasmic calcium
binding protein, and other prawn allergens (Kamat and others
2014). Chitin, the principal component of the crustacean shell
and its derivatives, can address allergic responses by enhancement
of innate immune system, alteration of Th1/Th2 balance forward
to Th1 cells, inhibition of IgE production, and suppression of mast
cell degranulation (Vo and others 2015).
Processing treatments, such as HHP, washing, cooking, and
others can significantly decrease heavy metals such as Cd con-
tent in Pacific oyster (Rasmussen and Morrissey 2007). It is in-
teresting to note that selenium present in shellfish is beneficial
because it can assist in elimination of mercury. Bjerregaard and
Christensen (2012) reported that selenite, seleno-cysteine, and
seleno-methionine, when administered via food, displayed dose-
dependent elimination of MeHg from marine shrimp. Analyses
of 485 samples of the 43 most frequently consumed shellfish and
fish species in Spain showed that these items possessed beneficial
Hg-to-Se ratios and Se-associated health benefit values (Olmedo
and others 2013).
A study on the dietary influence of cholesterol from shrimp
on levels of plasma lipoproteins (LDL and HDL) found that, al-
though consumption of steamed shrimp up to 300 g/d supplied
an amount of 590 mg dietary cholesterol, it did not impair the
lipoprotein profile. While the shrimp consumption increased LDL
by 7.1%, the beneficial HDL was increased by 12.1% in compar-
ison with a baseline diet. In contrast, consumption of 2 large
eggs per day resulted in an intake of 581 mg dietary cholesterol,
raising LDL by 10.2% (De Oliveira e Silva and others 1996). An-
other study reported that, while diets containing oyster, clam, or
crab lowered LDL tr iglycerides, and cholesterol, diet containing
shrimp did not change the ratio of LDL to HDL (Childs and
others 1990). Animal feeding studies reported a beneficial role of
hydrolyzed fish proteins in the reduction of plasma total choles-
terol together with increase blood HDL (Wergedahl and others
2004). In addition, noncholesterol sterols present in shellfish have
the potential to reduce the absorption of cholesterol (Dong 2001).
These studies suggested that intake of cholesterol from shellfish
is unlikely to adversely affect the consumer’s overall lipoprotein
profile. It may be mentioned that in the human body about two-
thirds of cholesterol is metabolically formed, while the remaining
is derived from foods. The cholesterol contents in shellfish can be
beneficial for normolipidemic populations for proper functioning
of the cell membranes and also as precursor for steroid hormones
and bile acids (Wardlaw and Smith 2009). The 2015 U.S. D ieta r y
Guidelines Advisory Committee observed absence of clear con-
nection between the intake of cholesterol and level of the sterol in
the blood. The Committee also found absence of adequate evi-
dence for a quantitative limit for dietary cholesterol specific to the
Dietary Guidelines. The high blood cholesterol level among con-
sumers could be linked to their intake of saturated fat, suggesting
a need to restrict intake of saturated fat (Anonymous 2015). It
may be further pointed out that shellfish have lower atherogenic
(a condition having inflamed plaques on the insides of arteries)
and thrombogenic (condition causing coagulation of blood) in-
dices (Bono and others 2012). An atherogenic index of 0.36 for
shrimp, in comparison with atherogenic indices of 1.0, 0.7, and
0.67 for mutton, beef, and pork, has been reported (Dayal and
others 2013).
The measures to ensure safety of farmed shellfish encompass
proper site selection, licensing, and certification of shellfish grow-
ing waters based on their hygienic quality, closure of farms in
cases of extreme contamination, regular monitoring of algae and
pathogens in the ponds, and proper sewage treatment. In addition,
control of stocking density, feed quality, shucking, and depura-
tion, and also introduction of a HACCP protocol will result in
improved quality of farmed shellfish (Patterson and others 1997;
Su and Liu 2013; Codex Alimentarius 2013). Rapid methods to
1234 ComprehensiveReviewsinFoodScienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®

Shellfish nutritive value and safety . . .
Table 10–Key areas to address shellfish associated hazards.
Classification, licensing, and certification of shellfish farms
Regular monitoring of environment and water quality
GMP implementation
HACCP implementation
Standard Operating Procedures (SOP)
Analytical testing: microbiological, physical, chemical, sensory.
Process verification and validation to ensure product safety
Traceability
Bar-coding of products
Labeling of products for consumer awareness
Import alerts, recall, and crisis management programs
Employee training and education
detect pathogenic microorganisms, biotoxins, allergens, heavy
metals, and antibiotics can significantly contribute to shellfish
safety (Fernandes and others 2015; FAO/WHO 2016; San-
tos and Ramos 2016). A multiplex polymerase chain reaction-
denaturing high-performance liquid chromatography (MPCR-
DHPLC) method has been developed for the rapid detection
of the pathogens V. cholerae, V. parahemolyticus, V. vulnificus, Vibrio
rmimicus, Vibrio alginolyticus,andL. monocytogenes in aquatic prod-
ucts (Zhan and others 2015). Bar-coding and phylogenetic analysis
can be used to detect multiple species of shellfish, such as crabs,
in commercial products (Haye and others 2012). Food traceability
(defined as the ease with which a product can be traced through-
out the supply chain) tracks down the history of food across the
entire supply chain, so that recalls of unsafe food items can be con-
ducted efficiently (Dandage and others 2017). Labeling products
can warn consumers of the health risks involved in consuming
raw or under cooked shellfish. Farmer awareness about the poten-
tial farming-related hazards, such as adverse effects of antibiotics,
offers significant scope to improve product safety (Swapna and
others 2012). It has been recognized that shellfish culture, shellfish
restoration, and nature conservation are related issues that need to
be addressed (Smaal and Wijsman 2010). Table 10 indicates the
key areas to control shellfish associated hazards.
Monitoring of food safety by regulatory bodies
In recent times, the importance of an integrated, multidisci-
plinary approach to food safety and quality throughout the en-
tire food chain has been recognized. International and national
regulatory bodies with defined rules and standards have estab-
lished appropriate control programs to ensure consumer safety
in fish trade. There is particular emphasis on the FAO’s strat-
egy to promote international harmonization and capacity build-
ing (Ababouch 2006). The WHO extends guidance to the bi-
valve shellfish industry to minimize the risks to human health
(WHO 2010,2015). The FAO, WHO, and other international
organizations provide scientific assessments on foodborne hazards
in compliance with international standards, guidelines, and rec-
ommendations promulgated in the Code of Practice for Live and
Raw Bivalve Mollusks (Codex Alimentarius 2013). The Global
Aquaculture Alliance (GAC) provides advice on best aquaculture
practices (Lee and Connelly 2006). The Fish Inspector, published
by FAO/INFOFISH regularly informs on global developments
with respect to seafood inspection, quality control, and technol-
ogy (www.infofish.org; accessed 2017 January 6). The NSA is an
international organization concerned with the biology, ecology,
production, economics, and management of shellfish resources
(National Shellfisheries Association, NSA 2017).
In the United States, agencies responsible for food safety in-
clude the FDA within the Dept. of Health and Human Services,
the Food Safety and Inspection Service (FSIS) and the Animal
and Plant Health Inspection Service (APHIS), within the Dept. of
Agriculture (USDA), and the NMFS, within the Dept. of Com-
merce. The FDA is responsible for protecting consumers against
impure, unsafe, and mislabeled foods. The Environmental Protec-
tion Agency (EPA) is involved in setting standards and tolerances
for pesticide residues in foods and feed. The EPA also provides
regular advisories and guidelines on shellfish and fish consump-
tion. Other agencies having food safety responsibilities include
the Centers for Disease Control and Prevention (CDC), and the
Natl. Institutes of Health (NIH) (Holley 2011). The Natl. Shellfish
Sanitation Program (NSSP) of the FDA provides guidance for the
sanitary control of molluscan shellfish intended for human con-
sumption (FDA 2015). A recently published guide provides to the
U.S. seafood industry information related to Food Safety Mod-
ernization Act, HACCP, and quality-related problems in seafood
(Zimmerman 2016).
In the EU, the EU-funded database assists seafood and aquacul-
ture industries to collate data on contaminants and to devise safety
measures (ECsafeSEAFOOD 2017). Imports of fish and seafood
are strictly regulated in the EU, requir ing health certification and
traceability (Seafish 2017). Key developments in EU fisheries pol-
icy and fish hygiene are regularly notified in the newsletter Fish-
Files Lite (Megapesca (Portugal) 2017). The U.K. Food Standards
Agency provides guidance to reduce the risk of vulnerable groups
to food hazards (U.K. FSA 2017). The Health Canada and the
Canadian Food Inspection Agency share food safety responsibility
in the country. In Australia and New Zealand, a comprehensive
food standards code is enforced by the state and territory gov-
ernments and also by food enforcement agencies (FSANZ 2017).
The Food Safety Law addresses the safety issues associated with
aquaculture in China (Broughton and Walker 2010).
Worldwide, shellfish safety is thoroughly ensured taking into
consideration the health guidance values for contaminants, stip-
ulated by regulatory agencies (Hellberg and others 2012; Khora
2014). The Joint Expert Committee on Food Additives (JECFA) of
the FAO/WHO at its 72nd meeting established a Provisional Tol-
erable Weekly Intake (PTWI) for inorganic mercury of 4 μg/kg
body weight (bw) (JECFA 2010). The PTWI values (μg/kg bw/d)
for MeHg, Cd, and Pb have been reported 0.23, 0.83, and 3.6,
respectively (Hellberg and others 2012). The U.S. FDA has set an
action level of 1.0 ppm for MeHg corresponding to an intake of
0.5 μg MeHg/kg bw/d (Hellberg and others 2012). Current Eu-
ropean standards regulate the levels of microbiological agents, phy-
cotoxins, and chemical contaminants in food (Gu´
eguen and oth-
ers 2011). The EFSA Panel on Contaminants in the Food Chain
(CONTAM) has established a tolerable weekly intake (TWI) of
4μg/kg/bw of inorganic mercury, 1.3 μg/kg/bw of MeHg
(EFSA 2012), and 2.5 μg/kg/bw of cadmium (EFSA 2011).
The Codex Standard for Live and Raw Bivalve Mollusks (Codex
Standard 292–2008) has limits for biotoxins including STX, OA,
DA, brevitoxin, and AZA groups (Codex Alimentarius 2013).
The maximum permitted levels of PSP toxins per killogram of
flesh of mollusk are as follows: STX group, ࣘ0.8 mg of STX
equivalent; OA group, <0.16 mg of OA equivalent; DA group,
<20 mg DA; BTX group, <200 mouse units or equivalent; and
AZA group, <0.16 mg AZA (FAO/WHO 2016). The EU Di-
rective 91/492/EEC permits maximum level of PSP toxin at 0.8
mg STX equivalent/kg shellfish and non-dl PCBs at 200 ng/g
w wt (EC 2009). Australia, New Zealand, Canada, and Japan
recommend 200 OA equivalent/kg shellfish (Khora 2014).
Recent surveys have shown that, in general, the levels of heavy
metals and chemical pollutants in shellfish species are too low to
C2017 Institute of Food Technologists®Vol.16,2017 rComprehensive Reviews in Food Science and Food Safety 1235

Shellfish nutritive value and safety . . .
pose serious hazards. About two-thirds of the total seafood sup-
ply, and 9 of the 11 most consumed shellfish (and also fish) in
the U.S. had low or very low Hg contents (Groth 2010). Aver-
age exposure of U.K. consumers to 24 metal elements includ-
ing Hg from fishery and other food items has generally declined
over time, and it has remained at levels too low to cause major
health concerns (Rose and others 2010). Analysis of 47 fishery
products by the U.K. FSA revealed that shellfish harbor very low
levels of environmental pollutants, far below the European reg-
ulatory limits (Seafish 2017). Regular consumption of mussel at
1 kg/wk for 26 wk by 102 healthy men and women aged 48 to 76
y did not cause concern regarding Hg, As, and Cd levels in their
blood (Outzen and others 2015). Average concentrations of heavy
metals in 6 commercial cephalopod species of South Korea were
in the order of Al >As >Cd >Pb >Hg. All the metal contents
were within the regulatory guidelines, and did not pose any threat
to consumers (Nho and others 2016). Detailed studies showed
that shellfish contamination from seawater offers a rather low risk
to the general French population (Gu´
eguen and others 2011).
Heavy metal contents in shellfish products in a popular seafood
market in India were within the maximum levels prescr ibed by
the EU and the FDA (Sivaperumal and others 2007). Storelli’s
group has conducted detailed surveys on heavy metal contamina-
tion of Mediterranean shellfish. They found that a 70-g serving
of mollusks resulted in intake of 0.89 μg/kg/bw of Cd, which
corresponded to 35.6% of PTWI of the metal (Storelli and others
2010). Bioaccessibility of heavy metals is an important factor in
determining their hazards, which has received much attention in
recent years. Gao and Wang (2014) observed that bioaccessibility
varies with metals. They found that silver from farmed oyster had
the lowest oral bioaccessibility (38.9% to 60.8%), whereas the val-
ues for Cu and Zn range from 72.3% to 93%; Cd and Pb had values
between those of Ag and Zn. Cano-Sancho and others (2015) re-
ported that arsenic from seafood was bioaccessible up to 89%,
while Hg was less than 50% bioaccessible. Fernandes and others
(2008) examined the chemical pollutants PCDD/F and PCB in the
most commonly consumed shellfish in Scotland including mussels,
oysters, and scallops. The estimated adult dietary intakes of these
pollutants arising from the consumption of a typical portion of
these foods in combination with an average U.K. diet were in the
range of 0.5 to 0.6 pg (WHO toxic equivalent, TEQ) (2005)/kg
bw/d. These values were within the tolerable daily intake of 2.0
pg (WHO toxic equivalent, TEQ) (2005)/kg bw/d, endorsed by
an Independent Expert Committee on Toxicology of Chemicals
in Food, Consumer Products and the Environment. Khora (2014)
has listed 16 countries that have regulatory programs to protect
public health from marine toxin contamination.
The presence of antibiotics in farmed seafood has declined since
the 1980s due to farmer awareness campaigns, regulatory con-
trol, and also to the introduction of vaccines against bacterial dis-
eases (Jennings and others 2016). The national chemical residue
monitoring programs that operate in the U.S. and Canada report
compliance with maximum residue level (MRL) (Holley 2011).
According to EU Directive 2003/89/EC, allergen labeling is a
requirement for all foodstuffs produced in the European Com-
munity (Edmonds 2006). Sulfite is currently listed in Annex III
Part B of Directive 95/2/EC as an authorized food additive and
is labeled E223 with a maximum permitted residue in crustacean
products set at 150 ppm (Edmonds 2006).
Local control authorities are involved in the classification of
mollusks and other shellfish growing areas, based on the possibil-
ity of environmental contaminations (Jennings and others 2016).
The 4 components of the U.S. FDA cooperative program include
classification of shellfish growing areas based on water quality, in-
spection to ensure sanitary measures, control of harvesting from
prohibited waters, and laboratory analysis (FDA 2017). The U.K.
Food Standard Agency provides guidance on the hygienic produc-
tion of shellfish (U.K. FSA 2017). Traceability of shellfish in supply
chains is essential to ensure product safety. The food traceabil-
ity regulations of 21 Organizations for Economic Co-Operation
and Development (OECD) countries provide practical guidance
on seafood authenticity (Charlebois and others 2014; Zhang and
Bhatt 2014). A recently launched cloud-based mobile applica-
tion is expected to increase the transparency of the supply chain
(Dandage and others 2017).
In addition to the control measures discussed above, consumer
alerts are also provided by regulatory authorities. Recent alerts
were related to the presence of PSP in oysters, V. cholerae in frozen
cooked prawns, and C. botulinum in canned seafood, which were
issued by the International Food Safety Authorities Network (IN-
FOSAN) developed by the WHO and the FAO (INFOSAN
2014/2015). Alerts were also issued by the U.S. FDA after de-
tection of the presence of drug residues from unapproved animal
drugs and/or unsafe food additives in imported aquaculture shrimp
and prawn (FDA 2016). Such alerts are also issued by the Food
Standard Agency (U.K. FSA 2017) and by the EU (Megapesca
(Portugal) 2017). These measures have significant roles to protect
the consumer from contaminated shellfish.
Risk-Benefit Analysis
The risks and benefits of shellfish consumption have been eval-
uated in several studies. An expert consultation under the lead of
FAO and WHO, which evaluated risks and benefits of shellfish
(and other seafood) concluded that consumption of seafood pro-
vided energy, protein, and a range of other important nutrients,
including the long-chain n-3 PUFAs. There is absence of probable
or convincing evidence of CHD associated with MeHg At levels
of maternal exposure to dioxins (from seafood and other dietary
sources) that do not exceed the Provisional Tolerable Monthly
Intake (PTMI) of 70 pg/kg/bw, established by the Joint Expert
Consultation; neurodevelopmental risk for the fetus is negligible
(FAO/WHO 2011; James 2013). Sirot and others (2012) reported
that consumption of approximately 50 g/wk of mollusks, crus-
taceans (and also lean fish) supplied the consumer the recom-
mended intake levels of n-3 PUFA, selenium, and iodine while
MeHg, Cd, dioxins, polychlorobiphenyls, Zn, Ca, and Cu re-
mained below the tolerable upper intake values. Olmedo and oth-
ers (2013) reported that the daily intakes of heavy metals from
43 most frequently consumed shellfish (and also fish) represented
very low percentages of their reference values, ranging from 0.1%
(Se) to 3.9% (Cu) for a person weighing 60 kg. Cardoso and
others (2010) observed that, although the consumption levels of
seafood (consisting of many species) by individuals var ied con-
siderably from 140 g/wk in the U.K. to 628.5 g/wk in Iceland,
the probability of exceeding the PTWI value for MeHg was low,
ranging from 0.04% in the U.K. to 9.61% in Iceland. A recently
developed app, “BeneFISHiary,” has been reported as useful to the
consumer to provide simultaneous information on the contents of
selenium, omega-3 fatty acids, and mercury in fishery products
(Pirckle 2016). Studies on risk-benefit analysis have concluded
that the benefits of nutrients grossly outweigh the risks among the
general population, when a variety of shellfish, whether marine
or freshwater origin, farmed or wild, is consumed (IOM 2007;
Hellberg and others 2012; Oehlenschl¨
ager 2012; Olmedo and
1236 ComprehensiveReviewsinFoodScienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®

Shellfish nutritive value and safety . . .
others 2013; Weichselbaum and others 2013; Domingo 2016).
The Second Intl. Conference on Nutrition (ICN2), held in Rome
in November 2014, confirmed the importance of shellfish and
other seafood as a source of nutrition and health (FAO 2016).
Conclusions
Shellfish items can satisfy the dietary requirements of many
nutrients. Their nutrient contents depend on species, habitats,
harvesting season, feed, and other factors. The health benefits de-
rivedfromshellfishalsodependonspecifictypeofthespecies
consumed, the frequencies of consumption, as well as quanti-
ties consumed. There is the potential for producing nutrition-
ally enriched farmed shellfish using feeds fortified with nutrients
such as long-chain PUFAs, vitamins, and carotenoids. Although
carotenoids have been commercially used to improve the color
of farmed shellfish, particularly shrimp, the nutritional value of
shellfish fed with astaxanthin, β-carotene, and other carotenoids
has yet to be explored. Shellfish items including abalone, mus-
sel, and snail possess compounds, which have interesting bioac-
tivities, such as immune-modulating, anti-inflammatory, antioxi-
dant, ACE-inhibitory, and other functions. There is the potential
for uses of these components for the development of functional
foods. The nutritional benefits of shellfish can be derived with-
out undue concerns of their safety. Shellfish-associated hazards are
manageable at the stages of harvesting, farming, processing, stor-
age, distribution, and consumption, with appropriate intervention
strategies and control measures by national and international regu-
latory agencies. With increasing globalization, international trade,
and rising consumer interests, a need persists for constant vigil to
ensure sustainable supply of a safe shellfish.
Acknowledgments
V.V. values the several inspir ing discussions he had with the
late Dr. D. S. Pradhan, Associate Director, Bio-Medical Group,
Bhabha Atomic Research Center, Mumbai, India. The article is
dedicated to Dr. Pradhan.
Author Contributions
The layout of the review was constructed by V.V. Both V.V. and
K.G. searched the literature. V.V. developed the manuscript. V.V.
and K.G. edited, and V.V. revised the final manuscript.
Conflict of Interest
The authors declare no conflict of interest.
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