1.

Introduction
Shrimp culture has expanded and intensified rapidly
around the world during the last two decades. The develop-
ment of nutritionally complete shrimp diets is necessary for
further intensification of this industry in years to come, and
knowledge of the shrimps nutritional requirements is essen-
tial to achieve this.
Shrimp, like all crustaceans, require dietary lipids, one
of the three main classes of organic nutrients, along with car-
bohydrates and proteins. Lipids represent the most concen-
trated source of energy of all, supplying approximately 9
kcal/g, about double of that contributed by either carbohy-
drate or protein (Mead et al., 1986). Research efforts are still
focusing on gaining a better understanding of the metabo-
lism of lipids in crustaceans, as well as on establishing
requirements for essential lipids throughout their life cycle.
This paper reviews the role of essential fatty acids and phos-
pholipids in shrimp nutrition, and published data concerning
their requirements. We also describe some available sources
of these nutrients with potential use in the formulation of
shrimp diets.
2. Essential fatty acids
Fatty acids (FA) are among the most common lipids;
they are long chain carboxylic acids that occur in many
diverse forms, with variations in the degree and kind of
branching, number of double bonds, presence of other func-
tional groups, and chain length (Kates, 1972). Most of the
natural saturated FA have unbranched structures with an
even number of carbon atoms and no double bonds.
Monoene or monounsaturated FA have in general an even
number of carbon atoms and one double bond. FAwith more
than one double bond are termed polyene or polyunsaturated
FA(PUFA); if more than three double bonds are present they
are generally known as highly unsaturated FA (HUFA). FA
may be designated by trivial or systematic names, or by the
shorthand nomenclature recommended by Holman (1966),
which designates chain length, number of double bonds and
the position of the double bond nearest to the terminal
methyl group, whose carbon is designated as the omega ()
or n carbon. Among PUFA and HUFA, one can recognize
the n-3 and n-6 families (Table 1).
Burr and Burr introduced the concept of essential fatty
acids (EFA) in 1929. The group of EFA is composed of
PUFAand HUFAof the n-3 and n-6 families, which are nec-
essary for normal physiological function in animals and
man. Their essentiality lies not only in their physiological
importance, but in the fact that, like many vitamins, they
cannot be synthesized de novo by the body, they must be
obtained either directly or as partially elaborated precursors
from the diet (Willis, 1987).
EFA play a major role in the biological function of the
cell membrane, not only in its formation and integrity, but
also in modulating membrane fluidity, enzymes, ion chan-
nels, and receptor properties. They also act as precursors in
the synthesis of hormone-like molecules known as
eicosanoids, a group of oxygenated products derived enzy-
matically from 20-carbon PUFA (Yehuda et al., 1997).
2.1. Linoleic acid
Linoleic acid (LOA) belongs to the n-6 family, and it is
a precursor for n-6 C20 and C22 polyenoic acids formed by
elongation and desaturation processes carried out by
enzymes (e.g., FA synthetases, desaturases). Oils rich in
linoleic acid include soybean, safflower, sunflower, walnut,
corn, cottonseed, pine nut, sesame, and groundnut (Padley et
al., 1994).
2.2. Arachidonic acid
Arachidonic acid (AA) is an important fatty acid of the
n-6 family. It can be produced from linoleic acid, its major
precursor, by alternating desaturation and elongation steps:
18:2n-6 18:3n-6 20:3n-6 20:4n-6. It is a major con-
stituent of membrane lipids and the principal precursor of
eicosanoids, which in turn may lead to the production of
prostaglandins (e.g., PGE2, PGF2, PGD2, PGI2),
Volume 2, Number 1 Priscilla Shirley, Editor-In-Chief
Summer 2004
THE ROLE OF ESSENTIAL FATTYACIDS AND PHOSPHOLIPIDS IN SHRIMP NUTRITION
Mayra L. Gonzlez-Flix
a
*, Addison L. Lawrence
b
, Delbert M. Gatlin, III
c
, Martin Perez-Velazquez
a
a
Departamento de Investigaciones Cientficas y Tecnolgicas, Universidad de Sonora, Rosales y
Nios Hroes s/n, A.P. 1819 C.P. 83000, Hermosillo, Sonora, Mxico.
b
TAES Shrimp Mariculture Project, Texas A&M University System, 1300 Port Street, Port Aransas, Texas 78373, USA
c
Department of Wildlife and Fisheries Sciences, 210 Nagle Hall, Texas A&M University, College Station, Texas, 77843-2258, USA.
*Corresponding author. Tel: +52-662-259-2169, Fax: +52-662-259-2197; E-mail: mgonzale@correom.uson.mx
Technical Bulletin
thromboxanes (e.g., thromboxane A2, thromboxane B2) and
leukotrienes (e.g., leukotriene B4, leukotriene C4).
Thromboxanes and prostaglandins are known to cause vaso-
constriction and platelet aggregation (Clandinin and
Jumpsen, 1997). Leukotrienes function in constriction of
bronchial airway musculature, vascular permeability, and in
interactions between endothelium and white blood cells.
These conjugated trienes are formed in response to immuno-
logic and nonimmunologic stimuli (Mayes, 1993). Thus
eicosanoids regulate many inflammatory and hypersensitivi-
ty reactions (Clandinin and Jumpsen, 1997). Eicosanoids
may also modulate the activity of ion pumps (Freeman et al.,
1990), K
+
and Ca
2+
ion channels (Schweitzer et al., 1990;
Schwartz et al., 1992), and neurotransmitter uptake and
release (Templeton, 1988).
AA can be obtained from dietary sources such as veg-
etable oil and meat; it is found in small quantities (0.5-1.5%)
in lard and tallow oils, liver meal, salmon oil, pollack liver
oil, cuttle-fish liver oil, short-necked clam oil, sardine oil,
skipjack oil, squid liver oil and herring oil (Tacon, 1987).
Though rare in the plant kingdom, it can be found in some
mosses and ferns, and is a major component of some marine
algae.
2.3. Linolenic acid
Linolenic acid (LNA) is the major fatty acid of plant
leaves, stems and roots. This essential fatty acid is the pre-
cursor of the n-3 family of HUFA (Klenk and Mohrhauer,
1960): 18:3n-3 18:4n-3 20:4n-3 20:5n-3 22:5n-
3 22:6n-3. The most readily available source of -
linolenic acid is linseed oil, which normally contains 45-
60% of this acid (Padley et al., 1994).
2.4. Eicosapentaenoic and Docosahexaenoic acids
Eicosapentaenoic acid (EPA) and docosahexaenoic acid
(DHA) are found in unicellular marine algae, brown
macroalgae, in moss cells and in many animal tissues (e.g.,
retina, brain). In general, marine fish, shrimp and mollusc
oils are rich dietary sources of these n-3 EFA; oils whose
EPA and DHA content constitutes over 20% of the total FA
present include cod liver oil, cuttlefish liver oil, short-necked
clam oil, sardine oil, skipjack oil, shrimp head oil and squid
liver oil (Tacon, 1987).
3. Fatty acid requirements in shrimp
Crustaceans have a limited ability for de novo synthesis
of the LOAand LNAfamilies of FA(Kanazawa et al., 1979a,
Kayama et al., 1980); consequently, they require a dietary
source of EFA. In early studies, Kanazawa et al. (1977, 1978,
1979b, c) demonstrated that Marsupenaeus japonicus
requires LOA, LNA, EPA and DHA as EFA, with the n-3
HUFA being the most essential, followed by LNA and by
LOA, respectively. Several authors have reported qualitative
requirements for EFA by different shrimp species like
Fenneropenaeus indicus (Read, 1981), F. chinensis (Xu et
al., 1993), Penaeus monodon (Catacutan, 1991),
Litopenaeus stylirostris (Leger et al., 1985), and L. vannamei
(Lim et al., 1997). However, quantitative requirements have
not often been reported. Kanazawa et al. (1979d) suggested
that a combination of EPAand DHAshould be included at an
optimum level of 1% of the diet for M. japonicus juveniles,
and later suggested that a dietary provision of 1% n-3 HUFA
could be considered as a minimal value for postlarval pe-
naeids (Kanazawa et al., 1979e). Shewbart and Mies (1973)
showed that optimum growth of Farfantepenaeus aztecus
was achieved by the addition of 1% LNA to the diet. In
growth experiments with P. monodon, Chen and Tsai (1986)
indicated a requirement for HUFA at 0.5-1% of the diet,
while Rees et al. (1994) observed that postlarvae can grow
well on an Artemia diet with n-3 HUFA ranging from 12 to
22 mg/g dry weight. Merican and Shim (1997) observed that
growth and survival responses of juvenile P. monodon
improved with a lower supplementation level of DHA rela-
tive to LNA. Interestingly, Glencross and Smith (1999)
reported that the magnitude of the growth response of P.
monodon was greater with LOAthan with LNA, and a simi-
lar effect was reported by Read (1981) in F. indicus. Xu et al.
(1994) suggested that for F. chinensis the requirement for
LNA may be between 0.7% and 1% of the diet, but once
DHA was adequately provided in the diet (around 1%),
growth, molt frequency and survival were significantly
greater than in animals fed a diet with 1% LNA. They con-
cluded that although n-6 FA like LOA and AA have benefi-
cial effects on growth and survival, n-3 FA, especially DHA
are the most potent EFA for this species. Lim et al. (1997)
evaluated the growth response and FA composition of juve-
nile L. vannamei fed different dietary lipids. They found that
menhaden oil, rich in n-3 HUFA, was better utilized by this
species, but among plant oils, those rich in LNAhad a high-
er nutritional value than those rich in LOA. They concluded
that both n-6 and n-3 FA appear to be essential in the diet,
although n-3 HUFA were required for maximum growth,
feed efficiency, and survival.
Because there are differences in the feeding habits, i.e.
from omnivorous to carnivorous, among penaeid species,
and even among developmental stages, generalizations con-
cerning requirements for FA should be made cautiously.
Kanazawa et al. (1979f) suggested that differences in the
effect of dietary LNA on growth of various shrimp species
may be partly explained by differences in the capacity for
bioconversion of LNA to HUFA. This capacity is extremely
low in M. japonicus, such that they are incapable of synthe-
sizing enough EPAand DHAfrom dietary LNAto meet their
requirements.
4. Phospholipids
Phospholipids (PL) are so called polar lipids, because
they possess at least one polar group. They are chemically
composed of di-acylated glycerol molecules with a phos-
phatic moiety attached to the C3 of the glycerol group. Their
fatty acid residues vary in chain length and degree of satura-
tion. According to Paltauff and Hermetter (1990), PL can be
divided into sphingophospholipids (e.g., sphingomyelin) and
glycerophospholipids, whose structure allows variation in
the nature of the alcohol esterified to the phosphate, leading
to different glycerophospholipid classes (Table 2).
4.1. Role of dietary phospholipids
PL are the major constituents of membranes and are
vital to the normal function of every cell and organ. They
maintain cell structure and function, and have regulatory
activities within the membrane and outside the cell. For
instance, they serve as second messengers in cell signaling,
an essential process in regulating cell growth, proliferation,
differentiation, metabolism, nutrient uptake, ion transport,
and even programmed cell death. In addition, there is evi-
dence that PL containing choline, sphingomyelin, and their
metabolites are important mediators and modulators of
transmembrane signaling (Zeisel, 1993).
PL act as emulsifiers and facilitate the digestion and
absorption of FA, bile salts and other lipid-soluble matters.
They also have a role in the transport of lipids, not only in
the transport of absorbed lipids from the gut epithelium into
the hemolymph, but also in the transport of lipids between
tissues and organs (Coutteau et al., 1997), given that PL are
constituents of lipoproteins. Phosphatidylcholine (PC) is
particularly important because it is an essential component
of these lipoproteins (Hertrampf, 1992). High density
lipoproteins (HDL) and very high density lipoproteins
(VHDL) are the main lipoproteins found in some crustacean
species (Lee and Puppione, 1978). PL also act as acyl donors
for the lecithin cholesterol acyltransferase (LCAT) to convert
cholesterol into cholesterol ester, but crustacean hemolymph
is likely to have a lower LCAT activity than mammalian
blood, since cholesterol esters are found only in trace
amounts (Mankura et al., 1980; Teshima, 1997).
Dietary PL may serve as a source of choline, inositol,
EFAor even energy, and for early stages of crustaceans it has
been suggested that PL present in the diet serve as a direct
source of these nutrients (Coutteau et al., 1997). Emphasis
has been given to the beneficial effect of PL for early stages
or juvenile shrimp because, even though some crustaceans
can synthesize PL (Shieh, 1969), their biosynthesis general-
ly cannot meet their metabolic demand (DAbramo et al.,
1981; Kanazawa et al., 1985).
4.2. Phospholipid composition
The effect of PL on growth and survival of crustaceans
seems to vary with the FAand the kind of compounds ester-
ified at the C-3 position with phosphoric acid. Apparently,
effective PL for M. japonicus need to possess choline and
inositol groups besides unsaturated FA, such as LOA, LNA,
EPA, and DHA. According to Kanazawa et al. (1985), PC
and phosphatidylinositol (PI) containing high levels of n-6
and n-3 FAprobably serve as the lipid moieties of HDLin M.
japonicus. DAbramo et al. (1982) suggested that crus-
taceans may also prefer dietary PL to triglycerides as a
source of EFA, and may even be precursors in the synthesis
of diglycerides and triglycerides (DAbramo et al., 1980).
4.3. Phospholipid requirements in shrimp
To date, several studies have demonstrated the benefi-
cial effect of supplementing PL to the diet of shrimp, such as
M. japonicus (Kanazawa et al., 1979g; Teshima et al., 1982;
Kanazawa et al., 1985; Teshima et al., 1986a, b; Camara et
al., 1997), F. chinensis (Kanazawa, 1993), P. monodon
(Piedad-Pascual, 1986), and L. vannamei (Coutteau et al.,
1996; Gong et al., 2000). In a study with M. japonicus,
Kanazawa et al. (1985) observed a possible interaction
between dietary soybean PC and n-3 HUFA. Growth and
survival rates increased with increasing levels of soybean
PC, and with increasing n-3 HUFA levels from 0% to 1%,
and decreased at 2% HUFA level when diets contained 3%
soybean PC; however, Kontara et al. (1997) failed to detect
any significant interaction between dietary soybean PC and
n-3 HUFA. Coutteau et al. (1996) reported that the growth
response of early postlarval L. vannamei was significantly
improved by addition of 1.5% soybean PC to the diet. With
increasing dietary level of soybean PC, higher proportions of
20:1n-9, EPA, and total n-6 PUFAwere observed in total FA
of shrimp tissue. Fatty acid composition of PC in shrimp tis-
sue was very much influenced by dietary levels of soybean
PC; increasing its level decreased the proportion of saturat-
ed FA while LOA and EPA increased.
4.4. Sources of phospholipids
All products of plant and animal origin contain PL, but
not all contain high levels. Products of plant origin which are
rich sources of PL are soybeans, sunflower seeds, rape seeds,
maize, and groundnuts. Excellent animal sources of PL are
egg yolk, brain, and eye tissues (Hertrampf, 1992).
Generally, fish eggs contain a large amount of PL. In Atlantic
herring (Clupea harengus), PL represent almost 70% of the
total lipids in the ripe eggs (Tocher et al., 1985), from which
PC accounts for 58%. Moreover, PL from neural tissues like
brain and eyes are rich sources of EFA. For instance, DHAis
normally found in higher concentrations in polar lipids
(Henderson and Tocher, 1987). Today soybean oil is the
main source of natural PL. Microorganisms such as bacteria,
algae, fungi, and yeast have been considered as prospective
sources of PL (Hertrampf, 1992).
5. Conclusions
Through experimental research we have learned that
n-3 HUFA are presumably required to achieve maximum
growth, feed efficiency, and survival of shrimp. However,
quantitative requirements for EFAby many species currently
cultured throughout the world have not been completely
defined. When studying dietary requirements for EFA,
emphasis should be made in the need for investigating inter-
actions among EFA, e.g., competitive interactions between
the n-3 and n-6 series of FA for -6 desaturase, or between
FAfor a given elongase. Because of these competitive inter-
actions among and between FA, the need for investigating
appropriate dietary ratios of FAarises, because feeding inad-
equate proportions of particular FA to shrimp may result in
biochemical imbalances and less fit animals. Another inter-
action that should be addressed is that between EFA and
other nutrients such as PL. Research using known PL
sources, e.g., purified PL of a particular class and known
composition of FA, will help understand their metabolic role,
and their importance in contributing to EFA of an organism.
A possible effect of total dietary lipid content on require-
ments of shrimp for EFA, or the consideration of an optimal
level of neutral (e.g., triglycerides) and polar (e.g., PL) lipids
in shrimp diets also should be addressed in further investiga-
tions. Those investigations can be pursued while searching
for alternative lipid sources to exploit in the aquaculture
industry.
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Common name # of C Family Shorthand nomenclature Systematic name Structural formula
Linoleic acid 18 n-6 18:2n-6 Cis, cis-9, 12- CH3(CH2)4[CH=CH-
octadienoic acid CH2]2 -(CH2)6COOH
-Linolenic acid 18 n-3 18:3n-3 all cis-9, 12, 15- CH3CH2[CH=CH-
octadecatrienoic acid CH2]3-(CH2)6COOH
Arachidonic acid 20 n-6 20:4n-6 all cis-5, 8, 11,14- CH3(CH2)4[CH=CH-
eicosatetraenoic acid CH2]4 -(CH2)2COOH
Eicosapentaenoic acid 20 n-3 20:5n-3 all cis-5, 8,11,14, 17- CH3CH2[CH=CH-
eicosapentaenoic acid CH2]5-(CH2)2COOH
Docosahexaenoic acid 22 n-3 22:6n-3 all cis-4,7,10,13,6,19- CH3CH2[CH=CH-
docosahexaenoic acid CH2]6-CH2COOH
Table 1. Some polyunsaturated and highly unsaturated fatty acids.
R-group Class Systematic name Abbreviation
-H Phosphatidic acid or 1,2-diacyl-sn-glycerol-3- PA
Phosphoric acid phosphoric acid
-CH2-CH2-N(CH3)3 Phosphatidylcholine 1,2-diacyl-sn-glycerol-3- PC
phosphorylcholine
-CH2-CH(NH2)COOH Phosphatidylserine 1,2-diacyl-sn-glycerol-3- PS
phosphorylserine
-CH2-CH2-NH2 Phosphatidylethanolamine 1,2-diacyl-sn-glycerol-3- PE
phosphorylethanolamine
Phosphatidylinositol 1,2-diacyl-sn-glycerol-3- PI
phosphorylinositol
Table 2. Principal glycerophospholipid classes: R-group chemical structure
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OH
OH
OH
OH
OH