Aquaculture Nutrition

2015

doi: 10.1111/anu.12266

..........................................................................................

Oceanography Department, Faculty of Science, Alexandria University, Alexandria, Egypt

Received 9 July 2014; accepted 22 October 2014

This study was conducted to evaluate the effects of dietary

taurine on growth performance and feed utilization of
Nile tilapia (Oreochromis niloticus) larvae. Four plant
protein-based, isonitrogenous (400 g kg
1 protein), isoenergetic (19 MJ kg
1) diets supplemented with four taurine
concentrations (0.0, 5.0, 10.0 and 15.0 g kg
1; designated
as T0, T0.5, T1 and T1.5, respectively) were prepared. The
diets were fed to triplicate groups of fish larvae (0.024 g
average body weight), to apparent satiation, three times
per day for 60 days. Larval growth rates and feed utilization efficiency were significantly improved with increasing
supplemental taurine up to 10 g kg
1 and decreased with
further taurine supplementation. The quadratic regression
analyses indicated that the maximum larval performance
occurred at about 9.7 g kg
1 of total dietary taurine. Fish
survival was significantly lower at 15 g kg
1 dietary taurine
than at other taurine levels. Body protein significantly
increased, while body moisture and ash decreased, with
increasing dietary taurine up to 10 g kg
1 and decreased
with further taurine supplementation to 15 g kg
1. Body
lipid was not significantly affected by dietary taurine concentration. A number of body amino acids (tryptophan,
arginine, histidine, leucine, isoleucine, valine, alanine, glycine, threonine and taurine) significantly increased with
increasing supplemental taurine up to 10 g kg
1 and then
decreased with further increase in dietary taurine levels.
The rest of body amino acids were not significantly affected
by dietary taurine. The present results suggest that about
9.7 g kg
1 dietary taurine is required for optimum performance of Nile tilapia larvae fed soybean meal-based diets.
KEY WORDS:

feed utilization, growth, larvae, Nile tilapia,

soybean meal, taurine

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2015 John Wiley & Sons Ltd

Correspondence: A.-F.M. El-Sayed, Oceanography Department, Faculty

of Science, Alexandria University, Moharram Bey 21511, Alexandria,
Egypt.
E-mail: abdelfatah.youssif@alexu.edu.eg

Tilapia culture has grown rapidly during the past two decades, so that tilapias are currently the second largest
farmed finfish group in the world, only after carps (FAO
2014). This rapid industrialization of tilapia production in
recent years has led to gradual shift in tilapia culture from
extensive and semi-intensive systems to more intensive
farming practices, with an increasing demand for quality
seeds and dependence on formulated feeds (El-Sayed 2006).
Therefore, the production of sufficient quantities of highquality seeds and the formulation of appropriate, costeffective feeds have become a major challenge facing tilapia
culture industry. This means that the profitability of tilapia
culture is directly related to the quality of the seeds used
and the quantity and quality of feed consumed by the fish.
The shortage of quality tilapia seed production to meet the
increasing farmers demand remains one of the major challenges facing the expansion of tilapia culture (El-Sayed 2006).
Therefore, considerable attention has been paid to larval rearing and nutrition of farmed tilapia during the past two decades. Similarly, the nutrient requirements and feeding
management of tilapia broodstock have been extensively studied (Gunasekera et al. 1996a,b; Gunasekera & Lam 1997;
El-Sayed et al. 2003, 2005; El-Sayed & Kawanna 2008).
The increasing demand for fish meal (FM) accompanied
by shortage in global supply has resulted in escalating FM
prices during the past few years (Tacon et al. 2012). Therefore, intensive efforts have been given to the replacement
of FM with less costly and more available plant protein
sources for aquaculture feed production. In this regard,

particular attention has been given to oil plant sources,

such as soybean meal (SBM), cotton seed meal, sunflower
meal and sesame seed meal, as a partial or total fishmeal
replacer in aquafeed industry (Tacon et al. 2011). Despite
that these sources have good protein contents and essential
amino acid (EAA) profiles, they are limited in a number of
EAAs, such as sulphur-containing amino acids (methionine
and cysteine) and lysine. They also contain many endogenous antinutrients including protease inhibitors, phytohaemagglutinin and antivitamins, which may negatively affect
their nutritional values (El-Sayed 2006).
Most of the ingredients of plant origin are also limited in
taurine (2-aminoethanesulfonic acid) which is an end product of metabolism of sulphur-containing amino acids. Taurine is often classified as amino acid, despite that it lacks a
carboxyl group. It is not also incorporated into protein
synthesis or degradation of mammalian tissues (Kuzmina
et al. 2010). However, taurine accounts for 3050% of the
entire amino acid pool, depending on the animal species
(Jacobsen & Smith 1968). Taurine is involved in many
physiological functions in mammals, including modulation
of immune response, calcium transport (Takahashi et al.
1992), retina development (Omura & Yoshimura 1999), bile
acid metabolism (Hofmann & Small 1967), osmotic regulation (Thurston et al. 1980) and endocrine functions (Kuzmina et al. 2010). It also plays an important role in the
development of both muscular and neural systems. Full
details of taurine synthesis and functions in fish and shrimp
are reviewed by El-Sayed (2014).
Taurine synthesis in fish varies widely among fish species,
depending on fish species and developmental stage, feeding
habits and feeding histories and the water environment in
which the fish lives. This could also be related to the variation in the activity of L-cysteinesulfinate decarboxylase
(CSD), which is a key enzyme for the oxidation and direct
conversion of cysteine to taurine or conversion of methionine into cysteine, mainly in the liver and brain (Jacobsen
& Smith 1968; Chang et al. 2013).
Although taurine is a non-essential nutrient, its inclusion in the diet could improve fish performance. For
example, marine fish species, such as Japanese flounder
(Paralichthys olivaceus), Red sea bream (Pagrus major)
and yellowtail (Seriola quinqueradiata), lack, or have low
ability of taurine synthesis due to the absence of or limited CSD activities (Goto et al. 2001; Yokoyama et al.
2001; Park et al. 2002; Takagi et al. 2005, 2008, 2011;
Kim et al. 2008). Dietary taurine supplementation may be
indispensible for these fishes, particularly if they are fed
plant-based diets.

On the other hand, studies on taurine synthesis and

physiological functions in freshwater fishes are contradictory. Some freshwater fishes, such as common carp, rainbow trout and Atlantic salmon, have been reported to
have the ability to synthesize taurine; thus, they may not
require exogenous supplemental taurine (Goto et al. 2001;
Yokoyama et al. 2001; Espe et al. 2008, 2012). In contrast,
taurine supplementation has been found essential for optimal performance of freshwater fish such as rainbow trout
(Gaylord et al. 2006, 2007), grass carp (Ctenopharymgodon
idellus) (Luo et al. 2006) and Nile tilapia (Goncalves et al.
2011). It is evident that taurine is conditionally essential
when these fishes are fed diets of plant origin and deficient
in methionine and/or cysteine. The essentiality of taurine
for freshwater fishes may also be affected by the feeding
habits and previous feeding histories of these fishes
(Gaylord et al. 2006).
The effects of dietary taurine supplementation on the
performance and biological functions of Nile tilapia
(Oreochromis niloticus) are not well understood. As far the
authors know, only one study investigated the response of
Nile tilapia larvae fed plant protein diets to supplemental
taurine (Goncalves et al. 2011). The preliminary results of
that study revealed that the larvae require 8 g kg
1 taurine
for optimum performance. However, the taurine range used
in that study was relatively narrow (28 g kg
1); and therefore, it is not known whether Nile tilapia larvae would
require higher dietary taurine levels. It is evident that more
research is urgently needed to study the effects of wider
exogenous taurine levels on the growth performance and
feed efficiency of different sizes and growth stages of Nile
tilapia fed protein sources of plant origins.
Therefore, this study was carried out at Oceanography
Department, Faculty of Science, Alexandria University,
Egypt, to investigate the effects of dietary taurine on growth,
feed efficiency, body composition and amino acid profiles of
Nile tilapia (O. niloticus) larvae fed soybean-based diets.

Newly hatched Nile tilapia (O. niloticus) larvae were

obtained from a private hatchery near Alexandria, Egypt.
The fish were stocked in a 1-m3 fibreglass tank filled with
dechlorinated tap water for 24 h for resting. Triplicate
groups of 200 larvae (0.024 g average weight) were stocked
in 140-L glass aquaria connected in a closed, recirculating
system containing a biological filter. The culture system

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Aquaculture Nutrition 2015 John Wiley & Sons Ltd

was also provided with continuous aeration using an air

compressor (BOYU; Boyu industries Co., Ltd., North City
Industrial Village, Raoping, China). Water temperature
was maintained at 27  1 C, while natural light was used
throughout the study. Faeces were siphoned each morning,
before the first feeding and about 10% of the water was
replaced with fresh dechlorinated water of the same temperature. Water quality parameters including dissolved oxygen (DO), ammonia (NH4N), nitrates (NO3N), nitrites
(NO2N) and pH were examined twice a week using
HACH test kit (Loveland, CO, USA). The average values
of these parameters throughout the study were as follows:
DO = 5.7  1.2 mg L
1,
pH = 7.8  0.10,
NH4
N = 0.081  0.002 mg L
1, NO3N = 0.72  1.61 mg L
1
and NO2N = 0.00 mg L
1.

Four SBM-based, isonitrogenous (400 g kg
1 cp), isoenergetic (19 MJ kg
1) diets were prepared, containing four
concentrations of taurine (0.0, 5.0, 10.0 and 15.0 g kg
1;
designated as T0, T0.5, T1 and T1.5, respectively). In fact,
when we started this series of experiments on taurine
requirement of Nile tilapia, we used five levels (0.0, 5.0,
10.0, 15.0 and 20.0 g kg
1) fed to fingerling fish (1.0 g).

We found that beyond 10.0 dietary taurine, the performance and survival of the fish were reduced substantially
(data are being processed for publication). Therefore, we
decided to reduce the inclusion levels to four (0.0, 5.0, 10.0
and 15.0 g kg
1) for broodstock study and larval study.
The composition and proximate analysis and amino acid
profiles of the diets are shown in Tables 1 & 2. The diets
were prepared as described by El-Sayed et al. (2013). The
fish were fed the test diets to apparent satiation, three times
per day (at 09.00, 13.00 and 17.00 h), for 60 days. The fish
in each aquarium were collected and weighed at 15-day
intervals, and the average weights were recorded. The
amounts of feed consumed by fish in each aquarium during
each feeding interval were also recorded.

At the termination of the study, all fish in each aquarium

were netted, counted, weighed to the nearest mg and stored
at
20 C for final body composition and amino acid
analyses. Initial body analyses were performed on a pooled
sample of fish, which was weighed and frozen before the
study. A sample of each test diet was also stored at

20 C for chemical analysis. Proximate analysis of moisture, protein, lipid and ash was performed according to

Table 1 Composition and proximate analysis (g kg
1 dry weight) of the test diets
Experimental diets
Ingredients

T0

T0.5

T1

T1.5

Fish meal
Soybean meal
Wheat bran
Taurine
Soybean oil
Fish oil
Vitamins and minerals mix1
Dicalcium phosphate
Binder (CMC)2
Total
Crude protein
Ether extract
Crude fibre
Ash
NFE3
Taurine
GE4

100
700
110
0.0
20
20
20
20
10
1000
404.0
81.0
31.3
141.0
342.7
0.9
18.78

100
700
105
5
20
20
20
20
10
1000
396.1
79.3
28.0
134.0
362.6
7.0
18.87

100
700
100
10
20
20
20
20
10
1000
398.8
82.1
35.0
128.0
356.1
11.0
18.93

100
700
95
15
20
20
20
20
10
1000
392.9
75.5
30.0
130.0
371.6
16.5
18.80

Vitamins & minerals mixture contains mg kg
1 or IU kg
1 of dry vitamins & minerals powder: Vit. A 2 200 000 IU., Vit. D3 1 100 000
I.U., Vit. E 1500 I.U., Vit. K 800 mg, Vit. B1 1100 mg, Vit. B2 200 mg, Vit. B6 2000 mg, Vit. H 15 mg, Vit. B12 4 mg, Vit. C 3000 mg, Iron
160 mg, Magnesium 334 mg, Copper 21.6 mg, Zink 21.6 mg, Selenium 25 mg, Cobalt 2.38 mg.
2
Carboxymethyl cellulose used as binder.
3
Nitrogen-free extract determined by difference.
4
Gross energy calculated based on 23.64, 39.54 and 17.57 KJ g
1 for protein, lipid and carbohydrate, respectively.
1

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Aquaculture Nutrition 2015 John Wiley & Sons Ltd

Table 2 Amino acid content (% dry weight) of the test diets

Protein productive value (PPV) 100 protein gain (g)=

protein fed (g) on dry weight basis
.

Experimental diets
Amino acid

T0

T0.5

T1

T1.5

Lysine
Methionine
Threonine
Tryptophan
Arginine
Phenylalanine
Histidine
Isoleucine
Leucine
Valine
Cysteine
Alanine
Glutamic acid
Glycine
Serine
Aspartic acid
Proline
Taurine

2.27
0.54
1.25
0.58
3.01
1.56
0.98
1.15
2.32
2.13
0.41
2.00
6.86
1.51
1.62
3.68
2.42
0.09

2.31
0.55
1.26
0.61
2.87
1.42
0.98
1.22
2.51
2.10
0.35
2.14
6.69
1.44
1.58
3.38
2.41
0.70

2.26
0.54
1.25
0.60
2.94
1.66
0.95
1.16
2.38
2.21
0.39
2.01
6.58
1.40
1.56
3.43
2.39
1.10

2.17
0.51
1.20
0.58
3.00
1.58
1.02
1.21
2.44
1.99
0.42
1.96
6.62
1.39
1.60
3.52
2.27
1.65

Simple linear and nonlinear regressions were performed to

correlate the relationships between fish performance and dietary taurine concentrations. Nonlinear and linear functions
were estimated by the least square method using the SPSS
program, version 12 (SPSS Inc., Chicago, IL, USA). All data
were also subjected to a one-way analysis of variance
(ANOVA) at a 95% confidence limit, using SPSS software.
Duncans multiple range test was used to compare means
when F-values from the ANOVA were significant (P < 0.05).

The present results showed that supplementation of dietary

taurine significantly affected (P < 0.05) the growth rates and
feed utilization efficiency of Nile tilapia larvae (Table 3).
Larval growth rates and feed utilization efficiency were significantly improved (P < 0.05) with increasing supplemental
taurine up to 10 g kg
1 and decreased with further taurine
supplementation. The quadratic regression analyses indicated that the maximum larval performance occurred at
9.7 g kg
1 of total dietary taurine. The equations representing the relationships between fish performance (y) and dietary taurine (x) were as follows:

standard AOAC (1995) methods. Amino acids profiles in

the diets and in the whole fish body (freeze dried) were
determined using an automated amino acid analyzer (Hitachi L-8500A; Hitachi, Ibaraki, Japan), as described by
Kim et al. (2005).

Growth rates and feed efficiency were calculated as follows:

Percentage weight gain (PWG) 100 Wf
Wi =Wi ;

PWG : y
36:261x2 707:82x 4001:9; R2 0:7887

SGR : y
0:0095x2 0:1836x 6:3169; R2 0:8215

Specific growth rate % SGR 100 ln Wf
ln Wi =t;

where Wi and Wf are initial and final weights (g), and t is
the time of experiment (days).

FCR : y 0:0049x2 0:0947x 1:5949; R2 0:7825

Larval survival was not significantly affected by taurine
supplementation up to 10 kg
1 (P > 0.05). Increasing

Feed conversion ratio (FCR) dry feed intake (g)=

fish live weight gain (g):

Table 3 Effects of dietary taurine supplementation on growth rates, feed utilization and survival (mean  SEM) of Nile tilapia fry
Experimental diets
Growth parameter

T0

1

Initial weight (g fish )

Final weight (g fish
1)
Percentage weight gain
Specific growth rate
Feed consumed (g fish
1)
Feed conversion ratio
Protein productive value
Survival (%)

0.024
1.18 
4817 
6.49 
1.82 
1.57 
23.32 
84.50 

T0.5
0.011d
48d
0.02d
0.087b
0.05b
0.85c
0.29a

T1

0.024
1.61 
6608 
7.01 
2.44 
1.54 
27.39 
86.33 

0.02b
87b
0.02b
0.050a
0.05b
1.24b
4.06a

0.024
1.94 
7997 
7.32 
2.68 
1.40 
35.55 
85.33 

T1.5
0.08a
337a
0.07a
0.017a
0.07a
2.24a
2.33a

0.024
1.46 
5983 
6.84 
2.43 
1.69 
26.22 
75.34 

0.03c
127c
0.04c
0.044a
0.02c
0.22b
1.45b

Values in the same row with different letters are significantly different at P = 0.05.

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Aquaculture Nutrition 2015 John Wiley & Sons Ltd

supplemental taurine to 15 kg
1 resulted in a sharp reduction in fish survival (P < 0.05).
Body composition of Nile tilapia larvae was significantly
affected (P < 0.05) by dietary taurine supplementation
(Table 4). Body moisture and ash decreased with increasing
dietary taurine up to 10 kg
1 and increased afterwards.
Body protein significantly increased with increasing dietary
taurine up to 10 kg
1 and levelled off with further taurine
supplementation to 15 kg
1. Body lipid was also significantly increased with increasing supplemental taurine up to
10 kg
1 and decreased with further taurine supplementation to 15 kg
1.
The following body amino acids (tryptophan, arginine,
histidine, leucine, isoleucine, threonine, valine, alanine, glycine and taurine) significantly increased (P < 0.05) with
increasing supplemental taurine up to 10 g kg
1 and then
decreased, or levelled off (leucine and taurine) with further
increase in taurine levels (Table 5). On the other hand,

other amino acids (lysine, methionine, phenylalanine, cysteine, glutamic acid, serine, aspartic acid and proline) were
not significantly affected by dietary taurine (P > 0.05).

Generally, marine fish and shrimp larvae lack the ability to

synthesize taurine from methionine through cysteinesulfinate decarboxylase (CSD) pathway (Brotons-Martinez
et al. 2004; Mayasari 2005). Therefore, they have been
reported to require exogenous taurine supplementation for
maximum development, growth, feed utilization and survival. For example, enriching live food such as Artemia
and rotifers with taurine improved morphology, development and performance of marine fish larvae (Salze et al.
2011; Yun et al. 2012). When larval red sea bream
(P. major) (Chen et al. 2004), European sea bass (Dicentrarchus labrax) (Brotons-Martinez et al. 2004), Japanese

Table 4 Body composition (g kg
1) (mean  SEM) on wet weight basis of Nile tilapia larvae fed the test diets
Experimental diets
Composition (g kg
1)
Moisture
Protein
Lipid
Ash

Initial

T0

692.00
187.89
40.69
85.62

740.22
147.82
44.85
66.35

T0.5





2.82
1.72a
0.36a
0.26b

729.41
166.00
51.11
60.87

T1





2.37
0.25b
1.19b
1.58a

T1.5

683.13
168.91
58.24
66.73






3.3
0.38c
1.96c
2.94b

703.20
168.07
54.69
72.21






1.56c
1.23c
1.86d
0.98c

Values in the same row with different letters are significantly different at P = 0.05.

Table 5 Amino acid profiles in whole body (mean  SEM) (% dry weight) of Nile tilapia fry fed the test diets
Experimental diets
Body amino acid

T0

Lysine
Methionine
Threonine
Tryptophan
Arginine
Phenylalanine
Histidine
Isoleucine
Leucine
Valine
Cysteine
Alanine
Glutamic acid
Glycine
Serine
Aspartic acid
Proline
Taurine
Total

3.66
1.47
1.87
0.54
2.46
1.79
1.22
2.64
3.29
2.43
0.77
2.93
6.57
2.60
1.74
5.58
3.10
0.13
44.83

T0.5




















0.04a
0.012a
0.08b
0.003d
0.05c
0.11a
0.017d
0.011b
0.02b
0.08b
0.011a
0.02b
0.34a
0.02c
0.07a
0.25a
0.20a
0.003c
1.06a

3.71
1.42
2.19
0.60
2.79
1.79
1.59
2.67
3.52
2.71
0.72
2.95
6.91
2.73
1.57
4.98
2.69
0.74
46.36

T1




















0.01a
0.08a
0.005a
0.005b
0.09b
0.066a
0.035b
0.02b
0.035a
0.11ab
0.026a
0.035b
0.29a
0.025b
0.005a
0.17a
0.16a
0.04b
0.84a

Values in the same row with different letters are significantly different at P = 0.05.

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Aquaculture Nutrition 2015 John Wiley & Sons Ltd

3.90
1.37
2.31
0.65
3.09
1.90
1.74
2.85
3.72
2.85
0.74
3.16
6.79
2.87
1.71
4.93
2.82
1.09
48.57

T1.5




















0.02a
0.13a
0.01a
0.007a
0.016a
0.05a
0.040a
0.051a
0.013a
0.027a
0.004a
0.05a
0.76a
0.045a
0.02a
0.03a
0.065a
0.09a
1.16a

3.63
1.42
1.91
0.58
2.60
1.83
1.29
2.70
3.59
2.65
0.78
2.91
6.32
2.66
1.76
5.06
3.05
1.15
45.84





















0.16a
0.09a
0.05b
0.001c
0.04bc
0.019a
0.004c
0.004b
0.13a
0.05ab
0.032a
0.048b
0.11a
0.015bc
0.22a
0.18a
0.19a
0.06a
0.52a

flounder (P. olivaceus) (Chen et al. 2005), California

yellowtail (Seriola lalandi) and white sea bass Atractoscion
nobilis (Rotman et al. 2012) were fed taurine-enriched
rotifers, larval growth, survival and body taurine were also
significantly improved. Supplementing microencapsulated
diets with taurine may also improve marine larval performance and survival (Takeuchi et al. 2001; Salze et al.
2012).
On the contrary, studies on the freshwater species rainbow trout (Yokoyama & Nakazoe 1992; Boonyoung et al.
2013), channel catfish (Robinson et al. 1978) and Atlantic
salmon (Salmo salar) (Espe et al. 2012) indicated that they
have the ability to synthesize taurine from CSD pathway.
Exogenous dietary taurine did not support the performance
and survival of these fishes. However, a number of other
studies indicated that some freshwater fishes may lack the
ability of taurine synthesis through CSD pathway, and, in
turn, they may require exogenous taurine for optimum performance and physiological functions. For example, taurine
supplementation (0.5%) was essential for optimal performance of juvenile rainbow trout fed soy protein concentrate-based diets (Gaylord et al. 2006, 2007). Taurine
supplementation also improved growth rates, feed digestibility and feed efficiency of carps (Liu et al. 2006; Luo
et al. 2006).
However, these studies were carried out on fingerling,
juvenile and grow-out stages, while the available information on the effects of dietary taurine on larval performance
of freshwater fishes, especially Nile tilapia larvae, is very
limited. In the present study, a taurine-free diet resulted in
poor growth performance, whereas 10 g kg
1 dietary taurine resulted in the best growth rates and feed efficiency.
However, the quadratic regression analyses indicated that
the maximum larval performance occurred at about
9.7 kg
1 of dietary taurine. This value is slightly higher
than that reported by Goncalves et al. (2011). But taurine
range used by Goncalves et al. (2011) was relatively narrow
(28 g kg
1), and the fish may have required higher taurine
levels if wider dietary taurine range had been used. This
result may indicate that Nile tilapia larvae are unable (or
have limited ability) to synthesize taurine from methionine
through CSD pathway, despite that methionine and cysteine in the test diets used in the present study were within
the range reported for optimum performance of Nile tilapia
(El-Saidy & Gaber 1998; Nguyen & Davis 2009; Furuya &
Furuya 2010). The low body taurine concentration in
the taurine-free group compared to those fed taurinesupplemented diets may also suggest that Nile tilapia larvae
did not receive sufficient taurine from the control diet, and

supplemental taurine was necessary. Similar results have

also been reported in white shrimp (Yue et al. 2013).
In the present study, dietary taurine at 9.7 g kg
1 level
was sufficient for optimum performance and biological functions, while further increase in taurine concentration lowered
larval performance. This suggests that when taurine was provided at higher concentrations, excessive taurine may have
been excreted to keep body taurine at optimum concentration. This process is energy-demanding, leading to increasing
energy consumption and therefore reducing or levelling off
growth performance (Yue et al. 2013). Similar findings were
reported in rainbow trout (Yokoyama & Nakazoe 1992) and
gilthead sea bream (Pinto et al. 2013). Excessive dietary taurine may also lead to cessation of growth rates through
reducing feed intake as has been reported in Japanese flounder (Park et al. 2002) and rainbow trout (Gaylord et al.
2006). Mayasari (2005) found also that excessive exogenous
taurine reduced moulting and survival of white shrimp
(Litopenaeus vannamei) larvae. The author referred that
result to the possible poisonous effect of taurine when
provided at excessive concentrations. This may explain the
increase of fish mortality in the present study with increasing
dietary taurine concentration beyond 10 kg
1.
Body protein in the present study was highest, while
body water and ash were lowest (P < 0.05) at 10 kg
1 dietary taurine. Further increase in dietary taurine led to a
decrease in body protein and an increase in both moisture
and ash contents. Similar results were reported on juvenile
turbot (Scophthalmus maximus) (Qi et al. 2012), presumably due to the stimulation effect of taurine on growth by
stimulating feeding (Carr 1982) and increasing protein synthesis and deposit when taurine was supplemented at optimum levels (Li et al. 2009).
In the present study, body taurine was significantly
increased with increasing dietary taurine supplementation
(P < 0.05). This means that body methionine was not used
for taurine synthesis, supporting the argument that Nile
tilapia larvae lack the ability to biosynthesize taurine and
indicating that supplemental taurine is necessary for their
optimum performance. As previously mentioned, marine
fish species, such as Japanese flounder (P. olivaceus), red
sea bream (P. major) and yellowtail (S. quinqueradiata),
also have low or negligible ability of taurine synthesis due
to the absence of or low CSD activities during intermediate
metabolism from methionine to hypotaurine (Goto et al.
2001; Yokoyama et al. 2001; Park et al. 2002; Kim et al.
2003, 2005, 2008; Takagi et al. 2005, 2006a,b, 2008, 2011).
Therefore, supplemental taurine may be indispensible, particularly if they are fed plant-based feed.

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Aquaculture Nutrition 2015 John Wiley & Sons Ltd

In conclusion, the present study suggests that Nile tilapia

larvae lack the ability to biosynthesize taurine from methionine through CSD pathway. However, more research is
needed to support this assumption. About 9.7 g kg
1 dietary taurine is required for optimum growth rates, feed efficiency and survival of these fish larvae.

The authors thank GISIS Company, Ecuador, for providing the taurine that was used in the present study.

Association of Official Analytical Chemists (AOAC) (1995) Official

Methods of Analysis, 16th edn. AOAC, Arlington, VA, USA.
Boonyoung, S., Haga, Y. & Satoh, S. (2013) Preliminary study
on effects of methionine hydroxyl analog and taurine supplementation in a soy protein concentrate based diet on the biological performance and amino acid composition of rainbow
trout [Oncorhynchus mykiss (Walbaum)]. Aquacult. Res., 44,
13391347.
Brotons-Martinez, J., Chatzifotis, S., Divanach, P. & Takeuchi, T.
(2004) Effect of dietary taurine supplementation on survival,
growth performance and feed selection of sea bass (Dicentrarchus
labrax) fry fed with demand-feeders. Fish. Sci., 70, 7479.
Carr, W.E.S. (1982) Chemical stimulation of feeding behaviour. In:
Chemoreception in Fishes (Hara, T.J. ed.), pp. 259273. Elsevier,
Amsterdam.
Chang, Y.C., Ding, S.T., Lee, Y.H., Wang, Y.C., Huang, M.F. &
Liu, I.H. (2013) Taurine homeostasis requires de novo synthesis
via cysteine sulfinic acid decarboxylase during zebrafish early
embryogenesis. Amino Acids, 44, 615629.
Chen, J.N., Takeuchi, T., Takahashi, T., Tomoda, T., Koiso, M.
& Kuwada, H. (2004) Effect of rotifers enriched with taurine on
growth and survival activity of red sea bream (Pagrus major) larvae. Nip. Sui. Gakk., 70, 542554.
Chen, J.N., Takeuchi, T., Takahashi, T., Tomoda, T., Koiso, M.
& Kuwada, H. (2005) Effect of rotifers enriched with taurine on
growth in larvae of Japanese flounder (Paralichthys olivaceus).
Nip. Sui. Gakk., 71, 342347.
El-Saidy, D.M.S.D. & Gaber, M.M.A. (1998) Amino acid requirements and composition of Nile tilapia (Oreochromis niloticus).
Ann. Agric. Sci. Moshtohor, 36, 163174.
El-Sayed, A.-F.M. (2006) Tilapia Culture, 274 pp. CABI Publishing, CABI International, Willingford, Oxfordshire, UK.
El-Sayed, A.-F.M. (2014) Is dietary taurine supplementation beneficial for farmed fish and shrimp, a comprehensive review. Rev.
Aquacult., 5, 115.
El-Sayed, A.-F.M. & Kawanna, M. (2008) Effects of dietary protein and energy levels on spawning performance of Nile tilapia
(Oreochromis niloticus) broodstock in a recycling system. Aquaculture, 280, 179184.
El-Sayed, A.-F.M., Mansour, C.R. & Ezzat, A.A. (2003) Effects of
dietary protein level on spawning performance of Nile tilapia
(Oreochromis niloticus) broodstock reared at different water
salinities. Aquaculture, 220, 619632.
El-Sayed, A.M., Mansour, C.R. & Ezzat, A.A. (2005) Effects of
dietary lipid source on spawning performance of Nile tilapia

..............................................................................................

Aquaculture Nutrition 2015 John Wiley & Sons Ltd

(Oreochromis niloticus) broodstock reared at different water

salinities. Aquaculture, 248, 187196.
El-Sayed, A.-F.M., ElGhobashy, A.E. & El-Mezayen, M.M. (2013)
Effects of feed colour on growth and feed utilization efficiency
of Nile tilapia (Oreochromis niloticus L.) fry and fingerlings.
Aquacult. Nutr., 19, 870876.
Espe, M., Hevry, E.H., Liaset, B., Lemme, A. & El-Mowafi, A.
(2008) Methionine intake affect hepatic sulphur metabolism in
Atlantic salmon (Salmo salar). Aquaculture, 274, 132141.
Espe, M., Ruohonen, K. & El-Mowafi, A. (2012) Effect of taurine
supplementation on the metabolism and body lipid to protein
ratio in juvenile Atlantic salmon (Salmo salar). Aquacult. Res.,
43, 349360.
FAO (Food and Agriculture Organization of the United Nations)
(2014) Global Aquaculture Production 19502012. FAO, Rome,
Italy
(http://www.fao.org/fishery/statistics/global-aquacultureproduction/en).
Furuya, W.M. & Furuya, V.R.B. (2010) Nutritional innovations
on amino acids supplementation in Nile tilapia diets. Rev. Bras.
Zootec., 39, 8894.
Gaylord, T.G., Teague, A.M. & Barrows, F.T. (2006) Taurine supplementation of all plant protein diets for rainbow trout (Oncorhynchus mykiss). J. World Aquacult. Soc., 37, 509517.
Gaylord, T.G., Barrows, F.T., Teague, A.M., Johansen, K.A.,
Overturf, K.E. & Shepherd, B. (2007) Supplementation of taurine and methionine to all plant protein diets for rainbow trout
(Oncorhynchus mykiss). Aquaculture, 269, 514524.
Goncalves, G.S., Ribeiro, M.J.P., Vidotti, R.M. & Sussel, F.R.
(2011) Taurine supplementation in diets for Nile tilapia (Oreochromis niloticus). World Aquaculture 2011, Natal, Brazil, 610
June, 2011. World Aquaculture Society, Abstract #639.
Goto, T., Tiba, K., Sakurada, Y. & Takagi, S. (2001) Determination of hepatic cysteinesulfinate decarboxylase activity in fish by
means of OPA-prelabeling and reverse-phase high performance
liquid chromatographic separation. Fish. Sci., 67, 553555.
Gunasekera, R.M. & Lam, T.J. (1997) Influence of dietary protein
level on ovarian recrudescence in Nile tilapia, Oreochromis niloticus (L.). Aquaculture, 149, 5769.
Gunasekera, R.M., Shim, K.F. & Lam, T.J. (1996a) Effect of dietary protein level on spawning performance and amino acid
composition of eggs of Nile tilapia, Oreochromis niloticus (L.).
Aquaculture, 146, 121134.
Gunasekera, R.M., Shim, K.F. & Lam, T.J. (1996b) Influence of
protein content of broodstock diets on larval quality and performance in Nile tilapia, Oreochromis niloticus (L.). Aquaculture,
146, 245259.
Hofmann, A.F. & Small, D.M. (1967) Detergent properties of bile
salts: correlation with physiological function. Annu. Rev. Med.,
18, 333376.
Jacobsen, J.G. & Smith, L.H. Jr (1968) Biochemistry and physiology of taurine and taurine derivatives. Physiol. Rev., 48, 424
511.
Kim, S.K., Takeuchi, T., Yokoyama, M. & Murata, Y. (2003)
Effect of dietary supplementation with taurine, b-alanine and
GABA on the growth of juvenile and fingerling Japanese flounder (Paralichthys olivaceus). Fish. Sci., 69, 242248.
Kim, S.K., Takeuchi, T., Akimoto, A., Furuita, H., Yamamoto,
T., Yokoyama, M. & Murata, Y. (2005) Effect of taurine supplemented practical diet on growth performance and taurine contents in whole body and tissues of juvenile Japanese flounder
(Paralichthys olivaceus). Fish. Sci., 71, 627632.
Kim, S.K., Matsunari, H., Takeuchi, T., Yokoyama, M., Furuita,
H., Murata, Y. & Goto, T. (2008) Comparison of taurine

biosynthesis ability between juveniles of Japanese flounder and

common carp. Amino Acids, 35, 161168.
Kuzmina, V.V., Gavrovskaya, L.K. & Ryzhova, O.V. (2010) Taurine Effect on exotrophia and metabolism in mammals and fish.
J. Evol. Biochem. Physiol., 46, 1927.
Li, P., Mai, K.S., Trushenski, J. & Wu, G.Y. (2009) New developments in fish amino acid nutrition: towards functional and environmentally oriented aquafeeds. Amino Acids, 7, 4353.
Liu, H., Li, H.W., Xu, Y.J., Shi, X.G. & Zhu, Z.C. (2006) Effects
of taurine on growth and nutritional value of carps. Food Sci.
Technol., 8, 097.
Luo, L., Wen, H., Wang, L., Li, Q., Long, Y., Guo, J.-L. & Yang,
X. (2006) Effects of taurine on growth performance, quality,
digestive and metabolic enzyme activity of grass carp
(Ctenopharymgodon idellus). Chin. J. Anim. Nutr., 18, 166171.
Mayasari, N. (2005) The effect of taurine to speed up molting and
increase physical endurance from vanname shrimp larva (Litopenaeus vannamei, Boone). World Aquaculture 2005, Bali, Indonesia, 9-13 May. World Aquaculture Society. Meeting abstract # 71.
Nguyen, T.N. & Davis, D.A. (2009) Methionine requirement in
practical diets of juvenile Nile tilapia, Oreochromis niloticus. J.
World Aquacult. Soc., 40, 410416.
Omura, Y. & Yoshimura, R. (1999) Immunocytochemical localization of taurine in the developing retina of the left eye flounder
(Paralichthys olivaceus). Arch. Histol. Cytol., 62, 441446.
Park, G.S., Takeuchi, T., Yokoyama, M. & Seikai, T. (2002) Optimal dietary taurine level for growth of juvenile Japanese flounder (Paralichthys olivaceus). Fish. Sci., 68, 824829.
Pinto, W., Figueira, L., Santos, A., Barr, Y., Helland, S., Dinis,
M.T. & Arag~
ao, C. (2013) Is dietary taurine supplementation
beneficial for gilthead seabream (Sparus aurata) larvae? Aquaculture, 15, 384387.
Qi, G., Ai, Q., Ma, K., Xu, W., Liufu, Z., Yun, B. & Zhou, H.
(2012) Effects of dietary taurine supplementation to a casein
based diet on growth performance and taurine distribution in
two sizes of juvenile turbot (Scophthalmus maximus L.). Aquaculture, 358359, 122128.
Robinson, E.H., Allen, O.W.J.R., Poe, W.E. & Wilson, R.P. (1978)
Utilization of dietary sulfur compounds by fingerling channel
catfish: L-methionine, DL-methionine, methionine hydroxy
analogue, taurine and inorganic sulfate. J. Nutr., 108, 19321936.
Rotman, F., Stuart, K. & Drawbridge, M. (2012) Effects of taurine
supplementation in live feeds on larval rearing performance of
California yellowtail (Seriola lalandi) and white seabass (Atractoscion nobilis). Aquaculture America 2012, Las Vegas, Nevada,
29 February-2 March, 2012. World Aquaculture Society.
Salze, G., Craig, S.R., Smith, B.H., Smith, E.P. & McLean, E.
(2011) Morphological development of larval cobia Rachycentron
canadum and the influence of dietary taurine supplementation. J.
Fish Biol., 78, 14701491.
Salze, G., McLean, E. & Craig, S.R. (2012) Dietary taurine
enhances growth and digestive enzyme activities in larval cobia.
Aquaculture, 362363, 4449.
Tacon, A.G.J., Hasan, M.R. & Metian, M. (2011) Demand and
Supply of Feed Ingredients for Farmed Fish and Crustaceans:
Trends and Prospects. FAO Fish. Aquacult. Tech. Paper No.
564, 87 pp. FAO, Rome.
Tacon, A.G.J., Hasan, M.R., Allan, G., El-Sayed, A.-F., Jackson,
A., Kaushik, S.J., Ng, W.-K., Suresh, V. & Viana, M.T. (2012)

Aquaculture feeds: addressing the long-term sustainability of the

sector. In: Farming the Waters for People and Food. Proceedings
of the Global Conference on Aquaculture 2010 (Subasinghe, R.P.,
Arthur, J.R., Bartley, D.M., De Silva, S.S., Halwart, M., Hishamunda, N., Mohan, C.V. & Sorgeloos, P. eds), pp. 193231.
2225 September 2010. Phuket, Thailand.
Takagi, S., Murata, H., Goto, T. et al. (2005) The green liver syndrome is caused by taurine deficiency in yellowtail, Seriola quinqueradiata fed diets without fishmeal. Aquacult. Sci., 53, 279
290.
Takagi, S., Murata, H., Goto, T., Hayashi, M., Hatate, H.,
Endo, M., Yamashita, H. & Ukawa, M. (2006a) Hemolytic
suppression roles of taurine in yellowtail (Seriola quinqueradiata) fed non fishmeal diet based on soybean protein. Fish.
Sci., 72, 546555.
Takagi, S., Murata, H., Goto, T., Ichiki, T., Endo, M., Hatate,
H., Yoshida, T., Sakai, T., Yamashita, H. & Ukawa, M. (2006b)
Efficacy of taurine supplementation for preventing green liver
syndrome and improving growth performance in yearling red sea
bream (Pagrus major) fed low fishmeal diet. Fish. Sci., 72, 1191
1199.
Takagi, S., Murata, H., Goto, T., Endo, M., Yamashita, H. &
Ukawa, M. (2008) Taurine is an essential nutrient for Yellowtail
(Seriola quinqueradiata) fed non-fish meal diets based on soy
protein concentrate. Aquaculture, 280, 198205.
Takagi, S., Hisashi, M., Goto, T., Hatate, H., Endo, M. &
Yamashita, H. (2011) Role of taurine deficiency in inducing
green liver symptom and effect of dietary taurine supplementation in improving growth in juvenile red sea bream (Pagrus
major) fed non-fishmeal diets based on soy protein concentrate.
Fish. Sci., 77, 235244.
Takahashi, K., Harada, H., Schaffer, S.W. & Azuma, J. (1992)
Effect of taurine on intracellular calcium dynamics of cultured
myocardial cells during the calcium paradox. Adv. Exp. Med.
Biol., 315, 153161.
Takeuchi, T., Park, G.-S., Seikai, T. & Yokoyama, M. (2001) Taurine content in Japanese flounder Paralichthys olivaceus T. & S.
and red sea bream Pagrus major T. & S. during the period of
seed production. Aquacult. Res., 32, 244248.
Thurston, J.H., Hauhart, R.E. & Dirgo, J.A. (1980) Taurine: a role
in osmotic regulation of mammalian brain and possible clinical
significance. Life Sci., 26, 15611568.
Yokoyama, M. & Nakazoe, J. (1992) Accumulation and excretion
of taurine in rainbow trout Oncorhynchus mykiss fed diets supplemented with methionine, cystine and taurine. Comp. Biochem.
Physiol., 102A, 565568.
Yokoyama, M., Takeuchi, T., Park, G.S. & Nakazoe, J. (2001)
Hepatic cysteinesulphinate decarboxylase activity in fish. Aquacult. Res., 32, 216220.
Yue, Y.R., Liu, Y.J., Tian, L.X., Gan, L., Yang, H.J., Liang, J.Y.
& He, J.Y. (2013) The effect of dietary taurine supplementation
on growth performance, feed utilization and taurine contents in
tissues of juvenile white shrimp (Litopenaeus vannamei, Boone
1931) fed with low-fishmeal diets. Aquacult. Res., 44, 13171325.
Yun, B., Ai, Q., Mai, K., Xu, W., Qi, G. & Luo, Y. (2012) Synergistic effects of dietary cholesterol and taurine on growth performance and cholesterol metabolism in juvenile turbot
(Scophthalmus maximus L.) fed high plant protein diets. Aquaculture, 324325, 8591.

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Aquaculture Nutrition 2015 John Wiley & Sons Ltd