Reviews in Aquaculture (2014) 6, 241–255 doi: 10.1111/raq.

12042

Is dietary taurine supplementation beneficial for farmed fish

and shrimp? a comprehensive review
Abdel-Fattah M. El-Sayed
Oceanography Department, Faculty of Science, Alexandria University, Alexandria, Egypt

Correspondence Abstract
Abdel-Fattah M. El-Sayed, Oceanography
Department, Faculty of Science, Alexandria Taurine is a neutral b-amino acid derived from the metabolism of sulphur-
University, Moharram Bey, 21511 Alexandria, containing amino acids. It is present in high concentrations in animal tissues,
Egypt. Email: afmelsayed@gmail.com especially heart, retina, skeletal muscle, brain, large intestines, plasma, blood cells
and leucocytes. Therefore, this amino acid plays significant roles in many physio-
Received 13 January 2013; accepted 20 June
logical functions, including membrane stabilization, antioxidation, detoxification,
2013.
modulation of immune response, calcium transport, myocardial contractility,
retina development, bile acid metabolism, osmotic regulation and endocrine
functions. Historically, taurine has not been considered as an essential nutrient
for fish. However, recent studies have indicated that taurine synthesis widely dif-
fers between fish species and demonstrated that it plays a key role in aquaculture
and nutrition of freshwater and marine fish and shrimp. Animal proteins are rich
in taurine, whereas plant proteins are taurine-deficient. Therefore, fish fed plant
protein–based food may require exogenous taurine for maintaining their physio-
logical functions. Nevertheless, taurine may be conditionally indispensable for fish
and shrimp, depending on dietary protein source, fish species and size, feeding
habits, previous histories and the rate of metabolism of its precursors, namely
cysteine and methionine. It is my belief that taurine functions in, and benefits for,
farmed fish and shrimp are now more than worthy of critical review and analysis.
This review summarizes the current knowledge on the roles of taurine in fish,
particularly farmed fish and shrimp, with emphasis on taurine structure and
biosynthesis, physiological functions, the effects of dietary taurine on fish perfor-
mance and health and live food enrichment with taurine.
Key words: fish and shrimp, metabolism, requirement, synthesis, taurine.

(Tacon et al. 2012). In the meantime, plant-based protein

Introduction
sources, particularly oil plant sources, have been receiving
Aquaculture continues to expand, with gradual shift from considerable attention over the past few decades as a partial
traditional, low-cost semi-intensive systems to more costly or total fishmeal replacer in aquafeed industry (Tacon et al.
intensive systems, where processed feed is a major compo- 2011). However, most ingredients of plant origin are lim-
nent. As a result, total industrial processed aquafeed pro- ited in a number of nutrients, including taurine, which
duction has increased from 7.6 million tonnes in 1995 to might be necessary for the optimal performance and
35 million tonnes in 2010, with an average annual growth metabolism of farmed aquatic animals, particularly for car-
rate of about 11%. It is also expected that this growth rate nivorous fish.
will continue, and aquafeed production will reach 70 mil- The beneficial effects of taurine on mammals have been
lion tonnes by 2020 (Tacon 2012; Tacon et al. 2012). The extensively investigated for several decades (Huxtable 1992;
aquaculture feed sector consumed over 73% of the total Stapleton & Bloomfield 1993; Di Leo et al. 2002; Lourencßo
global fishmeal production and over 80% of the total global & Camilo 2002; Kuzmina et al. 2010). However, only
fish oil production in 2010 (Tacon 2012). recently, the effects of taurine on fish physiology, metabo-
Fishmeals, which are the primary protein source in aqua- lism, culture and nutrition have been increasingly
feed industry, have become costly and of limited availability researched in different freshwater and marine fish species.

© 2013 Wiley Publishing Asia Pty Ltd 241

A.-F. M. El-Sayed

A good number of publications appeared on this subject route of taurine synthesis varies between species and depends
during the past two decades, with varying results. It is my on the type of tissue. For example, the rate of taurine synthe-
belief that comprehensive review and critical analysis of the sis in rats is higher than in human, whereas cats lack the abil-
role of taurine in aquaculture, and nutrition in particular, ity to synthesize this amino acid due to a limited activity of
appear timely. This review will tackle this issue, through cysteinsulphinate decarboxylase (CSD) (Knopf et al. 1978;
analysing and discussing the available studies that have Hayes & Trautwein 1989). CSD decarboxylates cysteinesulph-
been conducted on synthesis and functions of taurine in inate to hypotaurine, which, in turn, is oxidized to taurine
fish and its use in aquaculture and aquafeeds. It is hoped (Griffith 1987). Endogenous taurine synthesis occurs mainly
that this review will improve our understanding of taurine in the liver and brain in many mammalian species through
and its role in aquaculture. enzymatic oxidation and direct conversion of cysteine, or
conversion of methionine into cysteine (Jacobsen & Smith
1968; Stapleton et al. 1997; Stipanuk 2004; Chang et al.
Taurine structure and functions
2013). In rat liver, ‘taurine pathway’ accounts for 70% of deg-
Taurine, or 2-aminoethanesulfonic acid, is an end product radation of cysteine to pyruvate or taurine (Yamaguchi et al.
of metabolism of sulphur-containing amino acids. It is a 1973). In mammals, this process involves the actions of cyste-
neutral b-amino acid, where both the amine group and sul- ine dioxygenase (CDO), which leads to cysteinesulphinate,
phonic group can be ionized (Jacobsen & Smith 1968). and CSD (Griffith 1987). Pyridoxine (vitamin B6) is also
Taurine is not incorporated into proteins, nor degraded by required to convert cysteine or methionine to taurine
mammalian tissues (Kuzmina et al. 2010). However, it is (Brosnan & Brosnan 2006).
the most abundant free amino acid in animal tissue, Historically, taurine has not been considered as an essen-
because it accounts for 30–50% of the entire amino acid tial nutrient for fish (Robinson et al. 1978; Borlongan &
pool, depending on the animal species (Jacobsen & Smith Coloso 1993; Yokoyama & Nakazoe 1996; Akiyama et al.
1968). Taurine is concentrated mainly in the heart, retina, 1997). Just recently, studies have indicated that taurine syn-
skeletal muscle, brain, large intestines, plasma, blood cells thesis widely differs between fish species, depending on fish
and leucocytes in mammalian tissues (Huxtable 1992; species and size, feeding habits and cysteinesulphinate
Schuller-Levis & Park 2003). decarboxylase (CSD) activity. Marine fish species, such as
In mammals, taurine is involved in many biological Japanese flounder (Paralichthys olivaceus), red sea bream
functions, including antioxidation and detoxification (Sta- (Pagrus major) and yellowtail (Seriola quinqueradiata),
pleton & Bloomfield 1993), modulation of immune have low or negligible ability of taurine synthesis due to the
response, calcium transport (Takahashi et al. 1992), myo- absence of or low CSD activities during intermediate
cardial contractility, retina development (Omura & metabolism from methionine to cystathionine (Goto et al.
Yoshimura 1999), bile acid metabolism (Hofmann & Small 2001a; Yokoyama et al. 2001; Park et al. 2002; Kim et al.
1967), osmotic regulation (Thurston et al. 1980) and 2003, 2005a, 2008a; Takagi et al. 2005, 2006a,b, 2008,
endocrine functions (Huxtable 1992; Di Leo et al. 2002; 2011). Therefore, supplemental taurine may be indispensi-
Lourencßo & Camilo 2002; Kuzmina et al. 2010). This ble, particularly if they are fed plant-based food. On the
amino acid also plays a significant role in reducing the loss other hand, freshwater teleost fish, such as rainbow trout,
of potassium from heart muscles, and therefore can Atlantic salmon and common carp can synthesize taurine
prevent the onset of hazardous cardiac arrhythmias and through transsulphuration pathway (Goto et al. 2001a;
related heart disorders in humans (Eby & Halcomb 2006). Yokoyama et al. 2001; Espe et al. 2008, 2012; Kim et al.
Taurine has hypoglycaemic and antioxidant properties, 2008a). Tissue taurine concentrations of these fish
suppresses progressing of vascular complications in increased by dietary methionine and cystine supplementa-
humans and preserves secretory ability of pancreatic cells tion (Cowey et al. 1992; Yokoyama & Nakazoe 1992; Espe
(Hansen 2001). It also increases sensitivity of tissue recep- et al. 2008). Bluegill (Lepomis macrochirus) (Goto et al.
tors to insulin (Hansen 2001; Franconi et al. 2004). 2001a, 2004) also have the ability to synthesize taurine,
Taurine is also essential in the regulation and control of despite that the activity of CSD in bluegill fed a taurine-
serum cholesterol levels in the blood in humans (Lourencßo deficient diet was significantly lower than that of fish fed a
& Camilo 2002). high taurine diet (FM) (Goto et al. 2004). It appears from
these results that the natural feeding habits and feeding his-
tories of the fish may affect taurine biosynthesis, through
Taurine biosynthesis and metabolism
the effects on the activity of CSD (Gaylord et al. 2006).
Taurine biosynthesis is one of the most important catabolic This may explain the ability of carnivorous fish (such as
processes of sulphur-containing amino acids in most mam- Atlantic salmon, rainbow trout and channel catfish) and
malian species (Kuzmina et al. 2010). The predominant the inability (or low ability) of herbivorous fish (such as

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242 © 2013 Wiley Publishing Asia Pty Ltd
 Taurine requirement of farmed fish and shrimp

grass carp and tilapia) to synthesize taurine from methio- thus prevents cell death in animals and humans (Patrick
nine or cysteine. More discussion on the relationship 2006). Antioxidative and protective effects of taurine in fish
between taurine synthesis/requirement and feeding habits have recently attracted the attention of a number of
of the fish is provided later, under the heading ‘Effects of authors. Rosemberg et al. (2010) found that taurine treat-
taurine on the performance of farmed fish and shrimp’. A ment resulted in a significant reduction in lipid peroxides
full section ‘The relationship between fish size, feeding habits in zebrafish exposed to ethanol, due to the increase in total
and taurine requirement’ has been allocated to describing reduced thiol content detected after taurine pretreatment
this relationship. and alcohol exposure. Taurine has also been shown to alle-
viate cadmium (Cd)-induced oxidative stress in freshwater
catfish (C. batrachus) (Kumar et al. 2009). Similarly, tau-
Physiological functions of taurine in fish
rine was able to counteract oxidative stress induced by
As in mammals, taurine is involved in many physiological exposure to Cd in common carp (Cyprinus carpio) (Sevgiler
functions in fish, including osmotic regulation, antioxida- et al. 2011). Taurine can also protect body cells against oxi-
tion, feeding stimulation and retina development and dation, by protecting mitochondrial integrity and respira-
vision. This section briefly highlights the published works tion, as has been demonstrated in Atlantic salmon exposed
on these functions in different fish, crustacean and mollus- to CdCl2 (Espe & Holen 2013).
can species. Bile acids are steroids derived from cholesterol, synthe-
Because taurine comprises 30–>50% of the free amino sized by the liver, stored in the gall bladder and released
acid pool in blood, muscles and brains of animals, includ- into the intestinal lumen to emulsify fats and help in the
ing fish (Jacobsen & Smith 1968; Saha et al. 2000, 2002), it absorption of lipids and fat-soluble vitamins (Haslewood
may play a significant osmoregulatory role in the cellular 1967). Bile acids are conjugated mainly with taurine, and—
and central nervous system of marine fish species such as to a lesser extent—with glycine. Therefore, taurine may
flounder ((Pseudopleuronectes americanus) (King et al. enhance lipid metabolism in fish, through the increase in
1982) and skate (Raja erinacea) (Ballatori & Boyer 1992) as the activity of the bile salt-activated lipase in the liver, as
well as freshwater fish such as walking catfish (Clarias ba- has been reported in common dentex (Dentex dentex)
trachus) (Saha et al. 2000), tilapia and carps (Takeuchi (Chatzifotis et al. 2008). Bile salts (or free taurine) can also
et al. 2000, 2001a). For example, when walking catfish be used as olfactory stimuli for fish, and in turn, increase
levers were perfused with isotonic medium, their taurine feed consumption and/or change feeding behaviour (Sola
concentrations were lower than in fresh levers (Saha et al. & Tosi 1993; Chatzifotis et al. 2009). Marine fish such as
2000, 2002). The high physiological hepatic taurine concen- European glass eel Anguilla anguilla (Sola & Tosi 1993),
tration by the perfused liver suggests that taurine plays an European sea bass Dicentrarchus labrax fry (Brotons-Marti-
important role in cell volume homoeostasis in walking cat- nez et al. 2004) and gilthead sea bream Sparus aurata fry
fish (Goswami & Saha 2006). Similar findings have been (Chatzifotis et al. 2009) responded to taurine-supple-
reported in skate (Raja erinacea) (Ballatori & Boyer 1992) mented feeds more actively than taurine-free or taurine-
and trout (Oncorhynchus mykiss) (Michel et al. 1994). low feeds. Similar responses to dietary taurine have also
Takeuchi et al. (2000, 2001a) also found significant been reported in some freshwater fish. Both taurine- and
increase in taurine transporter mRNA in carp and tilapia glycine-conjugated bile acids were more stimulatory in
tissues during salinity-induced stress. The involvement of rainbow trout (Yamashita et al. 2006) and channel catfish
taurine in osmoregulation of crustacean and molluscan Ictalurus punctatus (Rolen & Caprio 2008) gustatory sys-
species has also been reported by many authors. Dalla Via tems. Feeding behaviour of juvenile Japanese flounder was
(1986, 1989) found that the most abundant individual also affected by dietary taurine (Kim et al. 2005b). Taurine
FAAs engaged in ionic regulation of juvenile penaeid shrimp supplementation (0.5 and 1.5%) led to the immersion of
(Penaeus japonicas) and prawn (Palaemon elegans) were the fish onto the bottom immediately after feeding, whereas
glycine, taurine, arginine, proline and alanine. Similar results the fish were swimming in the water column in the absence
have been reported in freshwater prawn (Macrobrachiumro- of taurine. The effects of dietary taurine levels and lipid
senbergii) (Tan & Chong 1981), where their transfer from contents on the conjugated bile acid composition of juve-
diluted to more concentrated media caused a significant nile Japanese flounder were further investigated by Kim
increase in FAA concentration in muscle tissue and a decrease et al. (2008b). When fish juveniles (0.04 g) were fed diets
in haemolymph protein content. Taurine has also been containing increasing concentrations of taurine (0.5–1.5%)
reported to accumulate in intracellular spaces of aquatic at two lipid levels (0 and 5%), body taurine contents
molluscs and is used as an osmolyte (Hosoi et al. 2008). increased with increasing dietary taurine. The conjugated
Taurine is also known to have antioxidant, membrane- bile acids in the gall bladder also increased with the increase
stabilizing properties, as it inhibits lipid peroxidation, and in the dietary taurine level and consisted of taurocholic acid

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A.-F. M. El-Sayed

(which accounted for more than 95% of the total conju- Lombardini 2002). Visual dysfunction in both human and
gated bile acids) and taurochenodeoxycholic acid. These animals may result from taurine deficiency (Militante &
results suggested that taurine is the sole precursor of conju- Lombardini 2002). This deficiency can be reversed with
gate bile acid in Japanese flounder juveniles. exogenous taurine supplementation. A number of studies
Taurine supplementation plays a significant role in pre- have been carried out on the role of taurine on retinal
venting green liver syndrome. This effect has been attrib- development and visual system in fish. It has been reported
uted to the increase in the excretion of bile pigments from that abundant taurine is localized in the retinal photorecep-
the liver into the bile and the decrease in the production of tor and neural layers of juvenile flounder (Omura &
haemolytic biliverdin (Takagi et al. 2005, 2006a). The Yoshimura 1999) and goldfish (Nusetti et al. 2009). These
effects of dietary taurine in the prevention of green liver results suggest that taurine may be involved in the protec-
syndrome have been demonstrated in red sea bream (Goto tion of the photoreceptor outer segment, the regulation of
et al. 2001b; Takagi et al. 2006b, 2010, 2011). When year- neural transmission and photoreceptor differentiation
ling fish were fed high SPC diets without taurine supple- during the developmental stages. In addition, Nusetti et al.
mentation, they exhibited inferior growth rates and feed (2009) found that taurine transporter, taurine and zinc also
efficiency, together with higher incidence of green liver coexist in photoreceptors and the ganglion cell layer,
(Takagi et al. 2006b, 2010). The hepatopancreatic taurine suggesting that both taurine and zinc are important in
concentration was also significantly lower, and hepatopan- normal cellular functions in fish retina.
creatic biliverdin concentration was higher compared with
the fish fed a FM-based diet. Taurine supplementation
Dietary sources of taurine
improved fish performance and alleviated these symptoms.
Taurine is an important neurochemical factor in the ani- Animal protein sources, such as meat, sea foods (fish, oys-
mal visual system (Omura & Yoshimura 1999; Militante & ters, mussels, clams, shrimp, crabs, etc.), eggs and dairy

1
Table 1 Taurine concentrations of foodstuffs (mg kg wet weight). Modified from Rana and Sanders (1986) and Stapleton et al. (1997)

Source Taurine Source Taurine

concentration concentration
(mg kg 1) (mg kg 1)

Meat Clams/fresh 2400*

Beef/raw 43*, 430† Clams/canned 1520*
Beef carcass 296† Scallop 8270†
Pork/raw 610* Squid 3560†
Chicken/raw dark meat 1690* Milk and derivatives
Chicken leg 337† Cow’s milk, homogenized 151†
Turkey/raw dark meat 3060* Yogurt/low fat plain 33*
Turkey 932† Ice cream vanilla 19*
Lamb/raw dark meat 470* Feedstuffs
Ham/baked 500* Meat and bone meal 956†
Seafood Meat meal 1150†
Tuna/whole 2840† Whey 660†
Tuna red meat 2798† Chicken viscera 1004†
Albacore tuna 1760‡ Poultry by-product meal 3049†
Tuna/canned 420* Fish scrap 1083†
White fish/raw 1510* Fishmeal 3201†
Wild salmon 600‡ Fish protein hydrolysate 7501†
Farmed salmon 600‡ Salmon meal 3106†
Greenland halibut 320‡ Shrimp meal 1094†
Plaice 1460‡ Tuna meal 1060†
Cod 1080‡ Yeast 112†
Mackerel 780‡ Phaeophyta (Laminaria japonica) 16.6†
Mussels/raw 6550* Rhodophyta (Gelidium subcostatum) 125†
Oysters/fresh 700*, 3960† Rhodophyta (Grateloupia elliptica) 24.8†
Cod/frozen 310*

*Lourencßo and Camilo (2002).

†Spitze et al. (2003).
‡Gormley et al. (2007).

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 Taurine requirement of farmed fish and shrimp

products, contain high levels of taurine (Jacobsen & Smith (Salmo salar) juveniles (Espe et al. 2012) and common carp
1968; Spitze et al. 2003). Marine algae also contain varying fry (Fontagne et al. 2000; Carvalho et al. 2004) and juveniles
levels of taurine. On the other hand, higher plant tissues (Kim et al. 2008a) indicated also that dietary taurine supple-
(such as soybean) are generally deficient in taurine. Taurine mentation is not necessary for optimum performance and
contents of the common food sources and feedstuffs are survival. These fish are able to synthesize taurine from
summarized in Table 1. methionine and cysteine through transsulphuration path-
way (Espe et al. 2008, 2012). Similar results were also
reported in juvenile red drum (Sciaenops ocellatus) (Patter-
Effects of taurine on the performance of farmed
son et al. 2012) reared in brackish water (7&). The nones-
fish and shrimp
sentiality of dietary taurine for these freshwater fish has been
In recent years, several publications appeared on the benefi- attributed to the transsulphuration of dietary methionine,
cial effects of supplemental dietary taurine on the growth, leading to an increase in body taurine in fish fed taurine-free
survival, feeding behaviour, feed consumption and feed uti- (or taurine-deficient) diets (Espe et al. 2008). The biosyn-
lization efficiency of different fish species and sizes. This thesis of taurine from methionine or cysteine has also been
section reviews and analyses these studies. For facilitating reported in adult rainbow trout (Yokoyama & Nakazoe
the issue, and assisting the reader extract the necessary 1991; Yokoyama et al. 2001). Taurine was accumulated in
information, the major results were piled in a comprehen- the whole body, liver and muscle when rainbow trout were
sive table (Table 2). fed with dietary sulphur amino acids, supporting the ability
of taurine biosynthesis in these fish. Subsequently, Yokoy-
ama et al. (2001) reported that the activity of cysteinesulphi-
Freshwater fish
nate decarboxylase (CSD) was high in rainbow trout.
As mentioned previously, exogenous taurine may not be In contrast to the above findings, a number of recent
essential for many freshwater fish species, because they have studies indicated that freshwater fish may require exoge-
the ability to synthesize it. Studies on taurine supplementa- nous taurine for optimum growth performance, feed con-
tion in farmed fish feeds indicated that supplemental methi- sumption, digestion and assimilation and other
onine and cystine with dietary taurine or inorganic sulphate physiological functions. This requirement depends on fish
have proved unsuccessful with fingerling channel catfish species, size and feed composition. For example, Gaylord
(Robinson et al. 1978) and rainbow trout (Page 1978). et al. (2006, 2007) reported that taurine supplementation
When these fish were fed dietary taurine, they exhibited (0.5%) was essential for optimal performance of juvenile
reduced growth and development of cataractogenesis in rainbow trout fed SPC-based diets (deficient in methio-
rainbow trout (Page 1978). Similarly, taurine was not bene- nine), while no effects of dietary taurine were observed in
ficial for fingerling and juvenile rainbow trout when they fish fed FM-based diets (rich in methionine and taurine).
were fed a methionine- or hydroxy analogue (MHA)-sup- Similarly, taurine supplementation improved growth rates,
plemented feed (Yokoyama & Nakazoe 1992; Boonyoung feed digestibility, feed efficiency, hepatopancreatic and
et al. 2012). When these fish were fed soy protein concen- intestinal protease, lipase and amylase activities of juvenile
trate (SPC) diet supplemented with MHA, they exhibited carp (species not provided) (Liu et al. 2006; Chunsheng
higher plasma taurine levels, presumably due to the conver- et al. 2007), grass carp (Ctenopharymgodon idellus) (Li
sion of dietary MHA into taurine, leading to promoted fish et al. 2005; Luo et al. 2006) and Nile tilapia (Oreochr-
growth (Boonyoung et al. 2012). These results suggest that omis niloticus) larvae (Goncßalves et al. 2011) fed plant pro-
dietary MHA may spare taurine in terms of promoting the tein–based diets. It is evident from these studies that taurine
performance of rainbow trout, while taurine had no effects is conditionally essential when these fish are fed diets con-
on fish fed both the SPC- and MHA-based diets. The final taining high plant protein levels and deficient in methionine
levels of taurine in the fish and fillet of rainbow trout juve- and/or cysteine. The essentiality of taurine for freshwater
niles fed plant proteins (full fat soy, extracted soy, soy pro- fish may also be affected by the feeding habits and previous
tein concentrate, corn gluten) which are deficient in taurine, feeding histories of these fish, as will be discussed later.
and fish hydrolosate (rich in taurine) were almost similar,
regardless of dietary taurine levels (Aksnes et al. 2006).
Marine fish and shrimp
Therefore, taurine seems to be homoeostatic regulated in
these fish independently of dietary taurine levels. This par- Much more efforts have been directed to the effects of tau-
ticular finding suggests that plant protein sources may be rine on cultured marine fish than freshwater fish. However,
included in diets for trout at high levels without affecting the most of the studies have been focusing on only few species,
feeding quality as evaluated by taurine levels. Studies on namely Japanese flounder (Paralichthys olivaceus), sparid
other freshwater species including Atlantic salmon fish (red sea bream Pagrus major and common dentex

Reviews in Aquaculture (2014) 6, 241–255

© 2013 Wiley Publishing Asia Pty Ltd 245
 246
Table 2 Dietary taurine levels tested and recommended for farmed fish and shrimp

Species Fish size (g) Taurine concentration (%) Remarks Reference

A.-F. M. El-Sayed

Tested (%) Required (%)

Freshwater fish
Rainbow trout 26.8 0.5–1.5 0.5 Taurine supplementation improved growth and feed utilization efficiency of fish fed the plant Gaylord et al. (2006)
(Oncorhynchus protein diets. No effects of dietary taurine were observed in fish fed the fishmeal-based diets
mykiss) 18.4 0.5 & 1.0 0.5 Dietary methionine was supplemented at 0, 0.5 and 1.0%. Taurine improved growth, while Gaylord et al. (2007)
methionine reduced growth. Taurine and methionine did not affect feed efficiency
Nile tilapia Newly hatched 0.2–0.8 0.8 Plant protein–based diets, growth increased with increasing taurine levels Goncßalves et al. (2011)
(Oreochromis niloticus) Larvae
Grass carp NA 0.02–0.18 0.06 Growth, feed efficiency, activities of lipase, amylase, transaminase and alkaline phosphatase Li et al. (2005)
(Ctenopharymgodon were significantly improved with taurine supplementation up to 0.06–0.8%
idellus)
Marine fish
Japanese flounder 0.9 0.5–1.5 1.4 White FM-based diets containing 0.5–1.5% of L-cystine. Taurine content in the diet affects the Park et al. (2002)
Paralichthys olivaceus sulphur amino acid metabolism
0.2 0.5 & 1.5 1.5 A combined mix of fish, krill and squid meal was the main dietary protein source. Growth rates Kim et al. (2005a)
and body taurine increased with the increase in dietary taurine level. The fish require at least
1.5% supplemental taurine
0.3 & 3.7 0.5 & 1.5 1.5 Ethanol-washed FM diets (taurine free). Performance improved with increasing taurine level. Kim et al. (2005b)
Abnormal feeding behaviour was observed at 0% taurine in the small-size fish (0.3 g)
0.7 0.1–1.5 0.5 Ethanol-washed FM diets (taurine free). Body taurine and conjugated bile acids contents Kim et al. (2007)
increased with the increase in dietary taurine.
9.6 0.1–1.5 1.5 Ethanol-washed FM diets (taurine free). Body taurine and conjugated bile acids contents Kim et al. (2007)
increased with the increase in dietary taurine
Summer flounder Juveniles 1.5 & 2.0 1.5 SBM-based diets. Up to 50–60% dietary SBM can be included in the diets without hindering fish Lightbourne (2011)
(Paralichthys dentatus) growth or survival when amino acids, phytase and at least 1.5% taurine were supplemented
Red sea bream 2.5 0.5–2.0 0.5–1.0 Semi-purified, casein-/gelatin-based diets. Dietary cysteine reduced fish performance. Body Matsunari et al. (2008a)
(Pagrus major) taurine content increased with the increase in dietary taurine.
4.7 0.1–0.7 0.5 Semi-purified casein-/gelatin-based diets. Growth, hepatic lipid levels and body taurine were Matsunari et al. (2008b)
increased with increasing dietary taurine
580.0 0.5–2.0 0.5 SPC-based diets. Taurine-deficient diet resulted in poor growth, incidence of green liver, low Takagi et al. (2010)
hepatopancreatic and plasma taurine and increase in hepatopancreatic bile pigment content
72.0 1.0 & 2.0 1.0 SPC-based diet. Taurine-deficient diet resulted in poor growth, incidence of green liver, low Takagi et al. (2011)
hepatopancreatic and plasma taurine and increase in hepatopancreatic bile pigment content
Turbot (Scophthalmus 6.3 0.5–1.5 1.0 A combined mix of casein, gelatin and FM (35, 11 and 10%, respectively) was the dietary Qi et al. (2012)
maximus) protein source. Fish performance improved with increased dietary taurine up to 1.0%, then
leveled off. The contents of taurine in body, muscle, eye, liver and brain were correlated with
consumed taurine

© 2013 Wiley Publishing Asia Pty Ltd

Reviews in Aquaculture (2014) 6, 241–255
 Table 2 (Continued)

Species Fish size (g) Taurine concentration (%) Remarks Reference

Tested (%) Required (%)

© 2013 Wiley Publishing Asia Pty Ltd

165.9 0.5–1.5 0.5 A combined mix of casein, gelatin and FM (35, 11 and 10%, respectively) was the dietary Qi et al. (2012)

Reviews in Aquaculture (2014) 6, 241–255

protein source. Fish performance improved with increased dietary taurine up to 0.5%, then
leveled off. The contents of taurine in body, muscle, eye, liver and brain were correlated with
consumed taurine
European Sea bass 0.81 0.1–0.3 0.2 FM/SBM-based diet. When fish were free to access the diets, they selected more actively the 0.2 Brotons-Martinez
(Dicentrarchus labrax) and 0.3% taurine-supplemented diets than the 0 and 0.1% taurine. When they were not et al. (2004)
allowed to choose, they exhibited better growth rate with diets containing 0.2 and 0.3%
taurine than those of 0 and 0.1% taurine, respectively
78 0.5–2.0% 1.0 High SBM, low FM protein source. The addition of 1 and 2% taurine promoted the growth and Kotzamanis et al. (2012)
increased muscle hardness and chewiness
Yellow tail 0.5 0.5- 2.0 1.0 A combined mix of fishmeal, krill meal and squid meal was the main dietary protein source. Matsunari et al. (2005a)
(Seriola Dietary taurine improved fish growth and affected the sulphur amino acid metabolism
quinqueradiata) 6100 0.5 & 1.0 1.0 A combined mix of FM, SBM and corn gluten meal (CGM) was the main dietary protein source. Matsunari et al. (2006).
The collection of eggs from females reared at 0% taurine was not successful. Oocyte growth
and spawning rates improved with increasing dietary taurine to 1%
Cobia 9.8 0.0 & 0.5 0.5 FM was partially replaced by yeast protein, with or without taurine, methionine or tryptophan. Lunger et al. (2007)
(Rachycentron At 50 and 75% yeast protein level, the best performance was recorded in fish fed diets
canadum) supplemented with all three amino acids or taurine alone
28 0 &0.5 0.5 Yeast protein–based diet supplemented with 0.5% taurine. Increasing yeast protein led to Lunger et al. (2007)
decreased weight gain and feed efficiency regardless of taurine supplementation
Marine shrimp
Grass shrimp (Penaeus NA 0.2–0.8 0.4 Purified diets. At 0.4, 0.6 and 0.8% taurine, performance was significantly better than at 0.0 or Shi-Yen and
monodon) 0.2% taurine Ben-Shan (1994)
White shrimp 0.48 0.04–0.2 0.17 Taurine supplementation significantly improved growth and feed efficiency compared with Yue et al. (2012)
(Litopenaeus taurine-free control diet
vannamei)

NA, not available.

Taurine requirement of farmed fish and shrimp

247
A.-F. M. El-Sayed

Dentex dentex), yellow tail (Seriola quinqueradiata), turbot logical functions of these fish when they are fed on plant
(Scophthalmus maximus), European sea bass (Dicentrar- protein–based diets. In support, early stage yellowtail juve-
chus labrax) and cobia (Rachycentron canadum), in addi- niles (0.5 g) fed on FM-based diet required only 1% taurine
tion to a few studies on marine shrimp. These studies for optimum growth performance (Matsunari et al. 2005a),
indicated that taurine may be conditionally essential for while larger juveniles (470 g) fed SPC-based diets required
these fish species. The effects of supplemental taurine on much higher dietary taurine (up to 4.5%) (Takagi et al.
the performances of these species depend on a number of 2006a, 2008). In addition, Takagi et al. (2005) reported that
factors including dietary protein sources, fish species and growth of juvenile yellowtail fed with SBM-based diets
size, natural feeding habits and previous feeding history of (58% soybean protein) was improved by 3% taurine sup-
the fish. plementation. Fish fed the SBM diet without taurine sup-
plementation resulted in inferior growth rates, low feed
efficiency, high mortality and anaemia, concomitant with
The relationship between dietary protein source and
higher incidence of green liver. This suggested that plant
taurine requirement
protein–based diets increase the requirement for taurine in
As mentioned earlier, plant protein sources (such as SBM) yellowtail juveniles, possibly to increase synthesis of tau-
are deficient in taurine or its precursors, namely cysteine rine-conjugated bile acids and to compensate for its excre-
and methionine (Espe et al. 2008). The inclusion of high tion from the intestine (Nguyen et al. 2011a,b). It is clear
levels of plant protein in marine fish feeds is expected to from this discussion that taurine is conditionally essential
cause deterioration in fish performance, possibly due to the for the investigated marine fish, when they are fed diets
inability, or low ability, of these fish to synthesize taurine. containing high concentrations of plant protein sources
For example, when red sea bream (P. major) were fed on (such as SBM) or diets deficient in sulphur amino acids. In
high plant protein diets (SBM, SPC), they were unable to the meantime, Lunger et al. (2007) found that when juve-
biosynthesize taurine from cystine (Goto et al. 2001b; Tak- nile cobia (Rachycentron canadum) fed diets containing 50
agi et al. 2006b, 2011; Matsunari et al. 2008a). Taurine and 75% yeast-based protein supplemented with taurine
supplementation (0.5–1%) was necessary for optimum alone, taurine, methionine and tryptophan or methionine
growth rates, feed efficiency, body taurine and physiological and tryptophan alone, the best performance was recorded
conditions. Supplemental taurine (0.2%) also improved in fish fed the yeast diets supplemented with all three
growth rates, feed efficiency and lipid metabolism of com- amino acids followed by the diets supplemented with tau-
mon dentex (Dentex dentex) juveniles fed SPC-based diets rine alone. The authors concluded that when taurine was
(Chatzifotis et al. 2008). The effects of taurine supplemen- supplemented alone, it allowed cobia to conserve the essen-
tation on growth performance of European sea bass (Dicen- tial amino acids (methionine and tryptophan), leading to
trarchus labrax) larvae (Brotons-Martinez et al. 2004) and improving growth rates.
juveniles (Kotzamanis et al. 2012) fed on SBM-based diets Processing of plant protein sources and taurine supple-
have been evaluated. Feeding activity and growth rates were mentation may also improve fish performance, feed diges-
improved with increasing dietary taurine supplementation. tion and synthesis of taurine-conjugated bile acids in
These results suggested that sea bass fry require 0.2% die- yellowtail (Nguyen et al. 2011a,b). Nguyen et al. (2011a)
tary taurine, while juvenile fish required 1% dietary taurine investigated the effects of SBM fermentation and taurine
for optimum performance. supplementation on lipid digestion and growth of finger-
Several trials have also been carried out to reduce the FM ling yellowtail fed SBM-based diets. Supplemental taurine
content in the diets of yellowtail, which is highly carnivo- or both taurine and fermentations improved lipid digest-
rous, by increasing the inclusion of plant protein sources ibility, lipase activity, lipid content and bile acid concentra-
(mainly soybean protein) in these diets. These diets have tion. However, growth rates were improved only by SBM
generally led to inferior performance and deterioration in fermentation with taurine supplementation. Similarly,
some physiological functions, mainly due to the deficiency growth rates, body taurine and bile acid levels of juvenile
of taurine (and other amino acids) in these plant sources, yellowtail fed diet containing digested and purified soy pro-
compared with fishmeals (Watanabe 2002; Takagi et al. tein isolate (DPSPI) as protein sources were better than
2008; Nguyen et al. 2011a,b). Taurine synthesis has been that of fish fed undigested, high molecular fraction (HMF)
reported to be extremely low or even negligible in yellowtail of soybean protein, soybean meal (SBM), soy protein iso-
(Takagi et al. 2005). The activity of CSD during intermedi- late (SPI), digested SPI (DSPI) (Nguyen et al. 2011b). In
ate metabolism from methionine to cystathionine is absent addition, taurine (1%), cholesterol (1%) and their combi-
or markedly low in these fish (Matsunari et al. 2005a). nation (1:1) resulted in a better growth rate of juvenile
Thus, it is expected that dietary taurine supplementation turbot fed high SBM diets than the control diet (Yun et al.
would be necessary for optimum performance and physio- 2012). Fish fed the taurine diet exhibited the highest

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248 © 2013 Wiley Publishing Asia Pty Ltd
 Taurine requirement of farmed fish and shrimp

activity of cholesterol 7a hydroxylase, followed by those fed utilization was not significantly enhanced. This suggests
the cholesterol and the control diet, respectively. that the increased growth may have been attributed to the
increase in feed intake, but not feed utilization efficiency.
Similar results have been reported in red sea bream (Matsu-
The relationship between fish size, feeding habits and
nari et al. 2008b; Takagi et al. 2011).
taurine requirement
It is evident from these studies that the response of mar-
In addition to the effects of protein source on taurine syn- ine fish to dietary taurine is affected by fish size (body
thesis, fish size has also been reported to affect taurine syn- weight). This phenomenon has been attributed to the tau-
thesis and metabolism in marine fish species. Extensive rine polyfunctionality appearing at different life stages of
research has recently been carried out on the effects of the animal (Kuzmina et al. 2010). The developmental stage,
exogenous dietary taurine supplementation on Japanese natural feeding habits and previous feeding histories of
flounder, at different growth stages, in terms of growth marine fish may therefore have a significant impact on tau-
rates (Park et al. 2002; Kim et al. 2005a,b), feeding behav- rine requirement and synthesis through the effects on the
iour (Kim et al. 2005b), conjugated bile acids contents activity of CSD (Gaylord et al. 2006). This supports the
(Kim et al. 2007, 2008b), sulphur amino acid metabolism assumption that taurine may be a conditionally essential
(Park et al. 2002; Kim et al. 2003) and taurine biosynthesis amino acid for larval and early juvenile stages of marine
(Kim et al. 2005a, 2008a). These studies indicated that Jap- fish such as Japanese flounder, turbot and yellowtail, which
anese flounder are unable to biosynthesize taurine from feed on taurine-rich foods during their early developmental
cystine, particularly during early stage juveniles (0.04– stages. However, Kim et al. (2007) reported that taurine
0.4 g). Supplemental taurine (1.5–2.0%) was necessary for supplementation improved the growth and feed efficiency
the improvement of growth, feeding behaviour, body tau- of both juveniles (0.7 g) and larger fingerlings (9.6) of Japa-
rine and sulphur amino acid metabolism, even if the fish nese flounder. The authors related this difference between
were fed with a taurine-rich protein sources (combined their results and the previous studies (Kim et al. 2003) to
mix of fish, krill and squid meal) (Park et al. 2002; Kim the dietary history of the fish before the start of the study.
et al. (2005a). This has been attributed the dependence of This particular result suggested that dietary taurine might
early stage flounder after settlement on taurine-rich mysids be an essential nutrient in both early and later life stages of
as the main natural food source (Hirota et al. 1990; Tanaka Japanese flounder.
1993; Furuta 1996; Sekai et al. 1997). On the other hand, Supplemental taurine has also been reported to improve
larger fingerlings (15 g) were not affected by taurine sup- reproductive performance of yellowtail broodstock (Matsu-
plementation (Kim et al. 2003). The growth rates and body nari et al. 2006). The fish were fed diets containing 40%
free taurine of turbot (S. maximus) larvae fed on natural FM and 24% SBM and supplemented with 0.0, 0.5 and
zooplankton or Artemia were also positively correlated with 1.0% taurine for 5 months prior to spawning. Oocyte
dietary taurine concentration, suggesting a dietary depen- growth improved significantly with the increase in dietary
dence of these larvae on taurine and/or sulphur amino taurine. Spawning success for females at these taurine con-
acids (Conceic
ao et al. 1997; Yun et al. 2012). In addition, centrations was 0.0, 14 and 86%, respectively. Fertilization
small juvenile turbot (6.3 g) fed casein-based diets required rate was also significantly increased, while eggs abnormality
higher dietary taurine (1.0%) for optimum performance, decreased, with increasing dietary taurine. The taurine con-
whereas larger fish (165.9 g) required only 0.5% taurine centrations in the liver and serum also increased with
(Qi et al. 2012). Matsunari et al. (2005a) found also that increasing taurine supplementation. This result suggested
supplementation of taurine (1%) to a FM-based diet fed to that supplemental taurine is an essential nutrient for
early stage yellowtail juveniles (0.5 g) for 6 weeks improved broodstock yellowtail, as it was found essential for larval
their growth performance over the first 3 weeks. After- and juvenile marine finfish, such as Japanese flounder and
wards, growth performance was not significantly related to red sea bream (Takeuchi et al. 2001b).
taurine supplementation.
Taurine supplementation has also been reported to
Marine shrimp
improve not only growth rates, but also feed consumption
and feed utilization of small fish sizes. Qi et al. (2012) Few trials have considered the effects of supplemental tau-
found that both feed intake and feed utilization and growth rine on physiological functions and growth performance of
rates increased in small turbot (6.3 g) by taurine supple- shrimps. These studies indicated that shrimp requires this
mentation. The authors suggested that increased growth amino acid for optimum performance during both larval
was probably due to the increase in feed intake and feed and grow-out stages. Shi-Yen and Ben-Shan (1994) found
utilization. In larger turbot (165.9 g), feed intake was sig- that grass shrimp (tiger shrimp) Penaeus monodon fed
nificantly increased by supplementation of taurine, but feed purified diets supplemented with 0.4–0.8% taurine had

Reviews in Aquaculture (2014) 6, 241–255

© 2013 Wiley Publishing Asia Pty Ltd 249
A.-F. M. El-Sayed

significantly higher growth and utilization efficiency and also resulted in higher enzyme activities for amylase, lipase,
serum cholesterol concentrations than at lower taurine trypsin and pepsin. In addition, when larval red sea bream
concentrations (0.0 and 0.2%). White shrimp (Litopena- P. major (Chen et al. 2004), Japanese flounder P. olivaceus
eus vannamei) fed low-fishmeal diets supplemented with (Chen et al. 2005), Pacific cod Gadus macrocephalus
taurine (0.04–0.4%) required 0.17% taurine for maximum (Matsunari et al. 2005b), California yellowtail and white
growth rates (Yue et al. 2012). Supplemental taurine (10 sea bass Atractoscion nobilis (Rotman et al. 2012) were fed
and 25 mg kg 1) also resulted in speeding up moulting taurine-enriched rotifers, larval growth, survival and body
rate of white shrimp larvae and significantly improved sur- taurine were significantly improved compared with taurine-
vival rate (Mayasari 2005). At higher taurine concentra- free rotifers.
tions (50 and 100 mg kg 1), all the larvae died, possibly Supplementing microencapsulated diets with taurine
due to the poisonous effect of taurine when provided at may also improve marine larval performance and survival.
excessive levels. However, further research is needed to sup- Takeuchi et al. (2001b) found that larval red sea bream
port this assumption. (P. major) fed with a microparticle diet supplemented with
Richard et al. (2011) evaluated the sparing effect of cho- taurine, together with rotifers, their body taurine was sig-
line and cystine on methionine requirements for protein nificantly increased. Pinto et al. (2010) also evaluated the
accretion in P. monodon juvenile (3
3 g) by measuring the effect of dietary taurine-supplemented microcapsules (dur-
activity of two key enzymes of remethylation (betaine– ing the pelagic phase) and artemia (during newly settled
homocysteine methyltransferase; BHMT) and transsulphu- phase) on Senegalese sole (Solea senegalensis) larvae. Tau-
ration (cystathionine b-synthase; CBS). They found that rine supplementation did not affect larval growth and
P. monodon has the ability to synthesize taurine, and this metamorphosis during the pelagic phase. However, sole
ability is significantly affected by the cystine level in the larvae previously fed microcapsules supplemented with tau-
30% SAA-limiting diets. The authors suggested that further rine had higher growth rate and metamorphosis success
research is needed to better understanding of the sparing than larvae fed control microcapsules. This particular find-
effects of both choline and cystine on methionine require- ing supports the assumption that previous feeding history
ments and also for characterizing the pathways regulating affects taurine synthesis and requirement in fish.
taurine synthesis in this species. On the contrary to the above results, which clearly indi-
cated the necessity of dietary taurine for larval development
of several marine fish species, dietary taurine supplementa-
Live food enrichment with taurine
tion does not seem to enhance the larval growth perfor-
It has been reported that rotifers Brachionus plicatilis and mance and survival of gilthead sea bream (Sparus aurata)
B. rotundiformis, which are indispensable live food organ- (Pinto et al. 2013). When the larvae were fed rotifers
isms for marine fish larvae, are deficient in taurine supplemented with taurine-enriched liposomes, growth
(Takahashi et al. 2005). Enriching rotifers with taurine performance, survival or larval taurine levels were not
may be necessary for improving larval rearing of marine significantly affected. When the larvae fed with live prey
fish. Therefore, enrichment of live food with exogenous were subsequently fed a solution containing L-methionine
taurine for marine fish larvae has been extensively investi- with or without supplemental taurine, dietary taurine sup-
gated and revealed that taurine is essential for larval plementation led to an increase in body methionine reten-
development and growth. Takahashi et al. (2005) found tion. This finding suggests that gilthead sea bream larvae
that the content of taurine in the enriched rotifers have the ability to synthesize taurine (through the conver-
increased with increasing taurine in the medium and sion of methionine into taurine). However, it would be
becomes constant after 16 h. The authors found also that unwise to build a conclusion on whether or not taurine is
enriched rotifers contained about 75% of initial accumu- essential for larval gilthead sea bream based on only one
lated taurine after 8 h from final enrichment. Salze et al. study. More work is urgently needed to support this
(2011) found that live rotifers and artemia enriched with finding.
4 g L 1 per day of taurine benefited larval morphology,
development (ontogeny of the lateral line system, olfac-
Conclusion
tory organ and gills), growth and survival of cobia
R. canadum. The same authors (Salze et al. 2012) found The present review showed that taurine synthesis widely
that cobia larvae fed either bioencapsulated diets or roti- differs between fish species, depending on fish species and
fer and artemia nauplii enriched with 4 g taurine L 1 per size, feeding habits and cysteinesulphinate decarboxylase
day exhibited improved development, growth, body tau- (CSD) activity. Generally, freshwater species have better
rine and survival rates and increased amylase and trypsin ability of taurine synthesis than marine species. However,
activities during early larval stages. Taurine supplementation some freshwater species have limited ability of taurine

Reviews in Aquaculture (2014) 6, 241–255

250 © 2013 Wiley Publishing Asia Pty Ltd
 Taurine requirement of farmed fish and shrimp

synthesis, and they require exogenous taurine source, espe- cysteine sulfinic acid decarboxylase during zebrafish early
cially if they are fed protein sources of plant origins. Marine embryogenesis. Amino Acids 44: 615–629.
fish species have low or negligible ability to synthesize tau- Chatzifotis S, Polemitou I, Divanach P, Antonopoulou E (2008)
rine due to a limited, or lack of, activity of CSD during Effect of dietary taurine supplementation on growth perfor-
intermediate metabolism from methionine to cystathio- mance and bile salt activated lipase activity of common den-
nine. Therefore, taurine is conditionally essential when tex, Dentex dentex, fed a fish meal/soy protein concentrate-
these fish are fed diets containing high plant protein levels based diet. Aquaculture 275: 201–208.
and deficient in methionine and/or cysteine, especially dur- Chatzifotis S, Arias MV, Papadakis IE, Divanach P (2009) Evalu-
ation of feed stimulants in diets for sea bream (Sparus aurata).
ing early life stages. Under such conditions, taurine supple-
The Israeli Journal of Aquaculture – Bamidgeh 61: 315–321.
mentation would be necessary for optimum growth rates,
Chen JN, Takeuchi T, Takahashi T, Tomoda T, Koiso M, Kuw-
feed efficiency, body taurine and physiological functions.
ada H (2004) Effect of rotifers enriched with taurine on
Live food or microcapsules enriched with taurine can also
growth and survival activity of red sea bream Pagrus major
improve larval morphology, development, growth and
larvae. Nippon Suisan Gakkaishi 70: 542–547.
survival of marine fish and shrimp. Taurine also plays Chen JN, Takeuchi T, Takahashi T, Tomoda T, Koiso M, Kuw-
significant osmoregulatory, simulative, protective and ada H (2005) Effect of rotifers enriched with taurine on
antioxidative roles in fish and shrimp, in addition to its growth in larvae of Japanese flounder Paralichthys olivaceus.
important role in retinal development and visual system. Nippon Suisan Gakkaishi 71: 342–347.
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