Amino Acid Requirements for Growth of Nile Tilapia
CORAZÓN B. SANTIAGO* ANDRICHARD T. LOVELLf
Binangonan Research Station, Southeast Asian Fisheries Development Center, *Binangonan, Rizal,
Philippines and ¿Departmentof Fisheries and Allied Aquacultures, Auburn university, Auburn, AL 36849
ABSTRACT A series of feeding experiments was con
ducted in aquaria to determine the quantitative require
ments of the 10 essential amino acids for growth of young
Nile tilapia (OreochromÃ-s nilotÃ-cus).The test diets con
tained casein and gelatin supplemented by crystalline L-
amino acids to provide an amino acid profile similar to 28%
whole egg protein except for the test amino acid. Each set
of test diets consisted of seven isonitrogenous diets con
taining varying levels of the amino acid to be tested. Weight
gains analyzed by the broken line regression method in
dicated the following requirements as a percentage of the
dietary protein: lysine, 5.12; arginine, 4.20; histidine, 1.72;
valine, 2.80; leucine, 3.39; isoleucine, 3.11; threonine, 3.75;
tryptophan, 1.00; methionine with cystine (0.54% of the
protein), 3.21; and phenylalanine with tyrosine (1.79% of
the protein), 5.54. J. Nutr. 118: 1540-1546, 1988.
INDEXING KEY WORDS:
•amino acids •h'He tilapia quantitative
requirements •fish nutrition
The indispensable amino acids for chinook salmon,
Oncorhynchus tshawytscha (I), sockeye salmon, O.
nerka (2), rainbow trout, Salmo gairdneri (3), channel
catfish, Ictalurus punctatus (4), common carp, Cypri-
nus carpio (5), red sea bream, Chrysophrys major (6),
two species of eel, Anguilla japónica and A. anguilla
(7, 8), and Zill's tilapia, Tilapia zilli (9) are the same 10
amino acids that are required by the rat. Quantitative
requirements for all of these amino acids have been
determined in growth trials for chinook salmon (10),
common carp, eel and channel catfish (11). The quan
titative requirements for rainbow trout (12) and Mos-
sambique tilapia (T. mossambicus] (13) have been es
timated from daily increases of essential amino acids
in body protein of fed fish, but without supportive growth
data.
Several species of tilapia (Tilapia and Oreochromis
spp.) are important culture fishes and knowledge of
their amino acid requirements would be useful in for
mulating research and commercial diets. The amino
0022-3166/88 $3.00 ©1988 American Institute of Nutrition. Received 13 fuly 1988. Accepted 30 August 1988.
1540
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acid requirements determined for the Mossambique ti
lapia by Jauncey, Tacón and Jackson (13) seem low in
comparison with those of other warmwater fishes, such
as channel catfish and common carp (11), and in com
parison with the essential amino acid percentages in
the muscle protein of the fish (13). The present study
was conducted to determine through growth trials the
quantitative requirements of the 10 indispensable amino
acids for growth of young Nile tilapia (O. niloticus}.
MATERIALS AND METHODS
Experimental diets. The basal diets were formulated
to contain 28% protein and 2.5 kcal of digestible ener
gy/g of diet, using purified ingredients. The dietary pro
tein level was below the optimum requirement for
growth of young Nile tilapia, 35% (14), to assure max
imum utilization of the limiting amino acid. Vitamin-
free casein and gelatin served as natural protein sources
for part of the protein and were supplemented with
crystalline L-amino acids to provide an amino acid pro
file similar to that of 28% whole egg protein except for
the test amino acid. Amino acid compositions of the
casein and gelatin were determined by a commercial
laboratory (Woodson-Tenent, Memphis, TN) and agreed
closely with published values (10). The amounts of ca
sein and gelatin in the basal diets were adjusted such
that the lowest level of the test amino acid came solely
from the intact proteins. Ingredient compositions of the
basal diets for the 10 tested amino acids are presented
in Table 1 and the crystalline amino acid supplements
for each test diet are presented in Table 2. The diets
containing varying levels of the test amino acid were
made isonitrogenous by decreasing the amounts of the
nonessential amino acids, aspartic acid, glutamic acid,
glycine and alanine, on a nitrogen basis, as the level of
test amino acid increased. The incremental levels used
for each test amino acid are given in Table 3. These
levels ranged such that the lowest and highest levels
were well below and above the anticipated require
ments, which were based on those of chinook salmon,
carp and channel catfish (10, 11).
 AMINO ACID REQUIREMENTS OF NILE TILAPIA 1541

TABLE1
Ingredient composition of basal diets used to determine requirements of the 10 essential amino acids for young Nile tilapia

Test amino acid

Ingredient Lys Arg His Thr Val Leu lie Met Phe Trp

CaseinGelatinDextrinCelluloseAmino 6.203.6027.3530.6719.7812.409.667.0223.4831.8815.5712.4010.974.2424.9

mixture1Other213.005.1523.4331.9714.0512.4014.371.3527.0230.5314.3312.407.195.4624.0132.0118.9312.404.482.0226.8131.2523.0412.406.993.3427.1130.7419.4212.40Ã
acid

'See Table 2 for amounts of crystalline L-amino acids used. Lysine and arginine were supplied as L-lysine • HC1 and L-arginine •HC1,
respectively.
2Amounts per 100 g dry diet: cod liver oil, 2.5 g; corn oil, 2.5 g; vitamin mix, 1.5 g; mineral mix, 4.0 g; and carboxymethyl cellulose, 1.9 g.
Vitamin mix provided the following diluted in cellulose (mg/kg diet]: thiamin, 20; riboflavin, 20; pyridoxine, 20; folie acid, 5; calcium
pantothenate, 200; choline chloride, 2,000; niacin, 150; ascorbic acid, 100; retinyl acetate (500,000 lu/g), 8; a-tocopherol (1 lu/mg), 50; chole-
calciferol (1,000,000 lu/g), 2; menadione sodium bisulfite, 10; inositol, 400; biotin, 2; cyanocobalamin, 0.06; ethoxyquin (antioxidant), 200.
Mineral mixture was that of Williams and Briggs (45) supplemented (in mg/kg diet) with CoCl2-6H2O, 4; A1K(SO4)2-12H2O, 5; and Na2SeO,,
0.2.

The diets were prepared as described by Wilson,
Harding and Garling (15) for channel catfish. The crys
talline amino acid mixture and the mineral mixture
were stirred into 375-430 ml of hot water (80°C). The
pH of the mixture was adjusted to 7.0 by gradually
adding 6 N NaOH. The other dry ingredients and the
oils were premixed and then blended with the water/
amino acid/mineral mixture. The moist mixture was
then passed through a food grinder to form 2-mm-di-
ameter moist pellets. The moist pellets were placed in
plastic bags and stored at - 10°C.Rations for 3-4 d
were thawed and stored in a refrigerator (5°C)
until fed.
The pellets were crumbled into smaller particles prior
to feeding. Test diets were freshly prepared for each
experiment.
Experimental design and feeding. Each experiment
consisted of seven diets containing different levels of
one test amino acid which were fed to fish in triplicate
aquaria. Diets were assigned to the aquaria in a com
pletely randomized manner. Twenty-one 60-1 aquaria
were supplied with constant-temperature (27 ±2°C) well
water with continuous aeration. Water exchange rate

TABLE2
Composition of amino acid mixtures added to the basal diets to provide an amino acid pattern similar to that of whole egg protein

acid
composition
L-Amino of 28% whole
acidsLysine1Arginine2HistidineThreonineValineLeucineIsoleucineMethionineCystinePhenylalanineTyrosineTryptophanGlycineAspartic egg protein1.921.820.731.461.8

acidGlutamic
acidSerineProlineAlanineLys00.9850.3970.8911.1571.3581.0430.8950.1080.9380.6890.25402.0350.8401.40600.851Arg0.85900.3890.9031.1681.3521.0310.9150.1040.9520

•Lysinewas supplied as L-lysine-HCl.

2Arginine was supplied as L-arginine-HC1.

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 1542SANTIAGO LOVELLTABLE
AND

3Percentage
growthyoung
weight gain, break point in forof curve and survival
levelsAmino Nile tilapia fed varying dietary 10acids'Mean
essential amino

of
replications/
fishDietary number of pointincrease
weight Break
acidLysineArginineHistidineThreonineValineLeucineIsoleucineMethionine3Number
replication%1.101.30•
level per amino%2335 level%±

1.432749
±892 0.082±
1493282±
1.401.501.601.701.900.600.801.001.201.401.601.800.200.300.400.500.600.700.800.200.400.600.801.001.201.600.400.600.800.901.001.201.400
2803287±
1962980
±
2762597
±
582929±
6835731
±
1.186445
±204 0.05±
1306574
±
626772±
±1116953
2486479
±
4255861
±
2821310
±
0.481485
±180 0.03,±
521672±
971712±
921462±
751653±
1221418
±
1081494
±
1.051730
±37 0.09±
2331924
±
492065±
1512405
±
542315±
1832493
±
2212191
±
0.782305
±180 0.05±
1232776
±
2242855
±
2442638
±
3672441
±
901495±
942832±
0.952138
±127 0.09±
3752137
±
2192492
±
3822565
±
1042735
±
4182215
±
1331484
±
0.871526
±187 0.08±
1831$57
±
4581658
±
1031652
±
1841953
±
1411837
±
811574±
0.751707
±168 0.06Meansurvival%818783909390839193
2561161
±
1211Õ97
±
2302000
±
1021929
±
5401 ±
87 ±618acid

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 AMINO ACID REQUIREMENTS
TABLE 3
ofreplications/number
fishDietary of
Amino replication0.60
level
acidPhenylalanine4TryptophanNumber per
3/1000.801.001.201.4051.601.800.05
3/500.100.150.200.250.350.45Initialweight1313131313131356565656565656Mean
'Regression of weight gain on dietary level of amino acid was significant at P < 0.05 for all amino acids.
2Standard deviation.
•'The
diet contained 0.15% cystine.
4The diet contained 0.5% tyrosine.
5Data from the 1.40% phenylalanine treatment were excluded from the regression analysis because of high disease-related mortalities in two
aquaria.
was once per 24 h. Measurements of oxygen and am
monia, using procedures described previously (14), in
dicated that the aquarium conditions were not stressful
to the fish. Walls and bottoms of aquaria were scrubbed
on approximately alternate days to minimize growth
of algae and fungi for the fish to feed on.
Fish used in the experiments were young Nile tilapia
(15-87 mg in size) produced at the Binangonan Re
search Station, Rizal, Philippines. The fish were stocked
randomly in the aquaria at rates of 45 to 100 per aquar
ium, depending upon size and availability of fry for the
experiment. Equal numbers of fry per aquarium were
used in each experiment. The fish were fed as much as
they would consume four times daily, at 0730, 1030,
1300 and 1600 h, for 8 wk.
The 10 experiments were conducted over an 18-month
period. At the end of each experiment, total weight and
number of fish in each aquarium were determined and
10 fish were randomly sampled from each aquarium
for total length measurement.
Statistical methods. Percentage gains in weight were
higher than percentage increases in length, so percent
age weight gain was used as the criterion for evaluating
growth response of the fish. The data were analyzed to
determine if there was a significant regression of weight
gain on increasing level of dietary amino acid, and if
the regressions were linear, quadratic or cubic (16). Where
significant nonlinear regressions of weight gain on amino
acid level were found, data were analyzed by the broken
line regression procedure of Robbins, Norton and Baker
(17). The broken line model determines the break point
in the growth curve which represents the optimum
dietary concentration of the amino acid for fish growth.
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OF NILE TILAPIA 1543
Continued
pointincrease
weight Break
level5242 amino acid
0.086520
±196 1.05 ±
7527476727846637902474918432082214121762409231174785440983527966
±
0.03138189391502572353 0.28 ±
285Meansurvival8889868740929097989897969797
RESULTS
There was a significant quadratic regression of growth
rate on dietary concentration of the test amino acid in
each experiment (P < 0.05). Survival rates were gen
erally high, ranging from 89% among all treatments in
the histidine experiment to 94% in the threonine ex
periment. In no experiment did mortality rate appear
to be diet related.
The young tilapia apparently utilized the test diets,
which contained up to 82% of the protein as crystalline
amino acids, relatively well. Maximum weight gain per
treatment ranged from 1672% increase of initial weight
for fish in the histidine experiment to 7902% increase
for fish in the phenylalanine experiment. Percentage
weight gains were higher for fish with lower initial
weights. The rates of gain were comparable to those
obtained in a companion study in this laboratory (18)
with fish from the same source and of similar size that
were fed natural aquatic food (mixed species of mi-
croalgae) and increased body weight by 2300 to 5600%
in 6 wk.
Lysine. Analysis of the growth data revealed a break
point in the growth response curve at 1.43% dietary
lysine (Table 3), indicating that increasing lysine be
yond this level would not provide significant additional
growth. This value corresponds to 5.12% of the dietary
protein (Table 4) and compares with 5.7% of dietary
protein for common carp (19), 4.3% for rainbow trout
(20), 5.0% for chinook salmon (21) and 5.0% for channel
catfish (22). This is much higher than the requirement
reported for the Mossambique tilapia of 3.78% of the
protein reported by Jauncey, Tacón and Jackson (13)
 1544 SANTIAGO AND LOVELL
and 4.05% reported by Jackson and Capper (23). No
gross anomalies were observed, such as caudal fin ero
sion in rainbow trout (20).
Arginine. The break point in the growth response
curve indicated that 1.18% dietary arginine provided
optimum growth from the experimental diets (Table
3). This corresponds to 4.20% of the dietary protein
(Table 4) and is similar to the values of 4.3, 4.3 and
4.5% for channel catfish (24), common carp (19) and
eel (11 ), respectively. It is lower than the requirement
of 5.8% determined for chinook salmon (25) and higher
than that of 3.5% determined for rainbow trout (12). It
is much higher than the value of 2.82% determined for
the Mossambique tilapia (13).
Excess arginine in the diets limited in lysine causes
depressed growth rate in chickens (26, 27) but not in
channel catfish (24). In this study, excess arginine caused
a reduction in growth rate (Table 3), however, there
was no lysine deficiency in the arginine test diets. In
fact, the lysine allowance in these diets exceeded the
determined lysine requirement by approximately 35%.
Histidine. The break point in the growth response
curve indicated that 0.48% histidine provided opti
mum growth for young Nile tilapia (Table 3). This cor
responds to 1.72% of the dietary protein (Table 4), which
is slightly below that of carp and eel, 2.1% (11, 19), but
near that of chinook salmon, 1.75% (25). The histidine
requirement of channel catfish, 1.54% of the protein
(28), is lower than that of Nile tilapia. The histidine
requirement reported for Mossambique tilapia (13),
1.05%, is much lower than that found for Nile tilapia.
Threonine. The threonine requirement was deter
mined to be 1.05% of the diet (Table 3) or 3.75% of the
protein (Table 4). As percentage of protein, the threo
nine requirement of Nile tilapia is near the requirement
reported for rainbow trout, 3.4% (12), carp, 3.9% (19)
and eel, 4.0% (11), but higher than the requirement
TABLE 4
Essential amino acid requirements for growth of young
Nile tilapia derived by the broken line method and presented
as a percentage of the protein
Amino acid Percentage of the protein
Lysine 5.12
Arginine 4.20
Histidine 1.72
Threonine 3.75
Valine 2.80
Leucine 3.39
Isoleucine 3.11
Methionine 2.68
Methionine plus cystine1 3.21
Phenylalanine 3.75
Phenylalanine plus tyrosine2 5.54
Tryptophan 1.00
'Cystine included as 0.54% of the protein.
2Tyrosine included as 1.79% of the protein.
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reported for channel catfish, 2.21% (29), chinook salmon,
2.25% (30) and Mossambique tilapia (13), 2.93%.
Valine. The break point in the response curve for
dietary valine showed the requirement to be 0.78%
(Table 3). This corresponds to 2.80% of the protein
(Table 4), which is lower than that reported for most
other fish. Valine requirement, as a percentage of the
protein, was 2.96% for channel catfish (28), 3.1% for
rainbow trout (12), 3.2% for chinook salmon (31), 3.6%
for common carp (19) and 4.0% for eel (11). The valine
requirement for the Mossambique tilapia as deter
mined by Jauncey et al. (13) was 2.2%. Exceeding the
dietary level of valine for optimum growth reduced
weight gain of Nile tilapia (Table 3).
Leucine. The leucine requirement was determined
to be 0.95% of the diet (Table 3) or 3.39% of the protein
(Table 4). This is about equal to that reported for chan
nel catfish, which is 3.3% (28), but generally lower than
values for other species, which are 3.9% for chinook
salmon (31), 4.4% for rainbow trout (12) and 5.3% for
eel (11). The leucine requirement for Nile tilapia agrees
closely with that determined for the Mossambique ti
lapia, 3.4% (13). Hughes, Rumsey and Nesheim (32)
reported the leucine requirement for lake trout (Sal-
velinus namaycush) to be 2.74 to 3.66% of the protein,
which may be comparable to that of the Nile tilapia.
Feeding a high level of leucine in the diet reduced weight
gain of Nile tilapia (Table 3).
Isoleucine. The break point in the isoleucine growth
response curve showed that the dietary requirement
was 0.87% (Table 3), or 3.11% of the protein (Table 4).
With the exception of eel, the isoleucine requirement
for which is reported to be 4.0% of the protein (11), the
requirement for Nile tilapia is generally higher than
that of other species. Channel catfish require 2.58%
(28), carp require 2.5% (12), chinook salmon require
2.2-2.7% (31) and rainbow trout require 2.4% (12).
Jauncey, Tacón and Jackson (13) reported 2.01% iso
leucine for the Mossambique tilapia.
Leucine-isoleucine interaction in Nile tilapia was not
apparent. Leucine level in the isoleucine test diets in
the present study was maintained at 2.55% of the diet,
which is the level in the reference amino acid profile
(28% whole egg protein). This is over two and one-half
times the leucine requirement of Nile tilapia. Isoleu
cine requirement of chinook salmon increased when
dietary leucine was in excess of the requirement (31).
Methionine. In the presence of 0.15% cystine in the
diet, which is the cystine level in the 28% whole egg
protein reference amino acid profile, the methionine
requirement was determined to be 0.75% of the diet
(Table 3) or 2.68% of the protein (Table 4). The total
sulfur amino acid requirement (methionine plus cys
tine) would be 0.90% of the diet, or 3.21% of the pro
tein. This agrees with the total sulfur amino acid re
quirements reported for chinook salmon, 3.75% (33),
carp, 3.1% (19) and eel, 3.2% (11). However, it is higher
than that reported for channel catfish, 2.34% (34), rain-
 AMINO ACID REQUIREMENTS
bow trout, 2.7% (12) and much higher than that esti
mated for Mossambique tilapia, 0.99% (13).
The replacement value of cystine for methionine is
from 56-66% for young pigs (35), from 48-53% for
chicks (36) about 50% for salmon (37) and 60% for
channel catfish (34) on a molar sulfur basis. The re
placement value of cystine for methionine in Nile ti
lapia has not been studied but can be presumed to be
approximately 50%.
Phenylalanine. In the presence of 0.5% tyrosine in
the diet, which is the level of tyrosine in the whole egg
protein reference amino acid profile, the phenylalanine
requirement is 1.05% of the diet (Table 3), or 3.75% of
the protein (Table 4). Total aromatic amino acid (phen
ylalanine and tyrosine) requirement is 5.54% of the
protein. This is similar to the total aromatic amino acid
requirements reported for carp, 6.0-6.5% (19) and eel,
5.8% (11), but higher than that for channel catfish, 5.0%
(38) and rainbow trout, 5.2% (12). It is much higher
than the total aromatic amino acid requirement deter
mined by Jauncey, Tacón and Jackson (13) for Mossam
bique tilapia, 2.5%.
About 50% of the total aromatic amino acid require
ment in channel catfish could be supplied by tyrosine,
and high levels of tyrosine (10% of the diet) had no
adverse effect on growth of the fish (38). Tyrosine has
been found to supply 42, 45 and 49% of the total aro
matic amino acid needs of young chicks (39), growing
rats (40) and weanling pigs (41), respectively. Rate of
phenylalanine replacement by tyrosine in Nile tilapia
is not known but can be assumed to be about 40 to
50%.
Tryptophan. The break point in the tryptophan growth
response curve showed the dietary requirement to be
0.28% (Table 3), or 1.00% of the protein (Table 4). This
is near the tryptophan requirement of eel, 1.1% of the
protein (11), but higher than that reported for most
other fish. Tryptophan requirement for maximum
growth of rainbow trout was reported to be 0.45% (42)
to 0.58% (43), for chinook salmon, 0.4-0.6% (44), chan
nel catfish, 0.5% (29) and common carp, 0.8% (19). The
tryptophan requirement for Mossambique tilapia was
reported to be 0.43% of the protein (13). No gross mor
phological signs of tryptophan deficiency, such as sco-
liosis, fin erosion and cataracts (43), were observed in
the Nile tilapia fed the tryptophan deficient diets.
DISCUSSION
The quantitative essential amino acid requirements
for growth of young Nile tilapia have been determined.
Favorable growth rates for the fish in this experiment
indicate that the diets were satisfactory. Although gen
erally similar to the requirements reported for other
fish species, amino acid requirements of Nile tilapia do
not completely agree with the requirements reported
for any one species. No requirement was excessively
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OF NILE TILAPIA 1545
high or low when compared with the range in require
ments determined for the other species. No overt de
ficiency signs were observed for any amino acid.
The amino acid requirements reported for the Mos
sambique tilapia, on a protein percentage basis (13), are
decidedly lower than those found here for the Nile ti
lapia. It is unlikely that the requirements differ this
much between these species. The requirements for the
Mossambique tilapia, which were not based on growth
data, were projected for a 40% protein diet; however,
when presented as a percentage of the diet, they are in
more agreement with those found for the Nile tilapia.
Because of the great variation in daily protein con
sumption by fish, caused by variations in size, feeding
practices and diet formulations, amino acid require
ments should be reported as a percentage of the dietary
protein, as in Table 4, rather than as a percentage of
the diet.
The amino acid requirements presented in Table 4
and in the National Research Council tables for other
species generally represent near maximum growth rate.
For practical feeding, these values may not be opti
mum. The procedures used to evaluate the growth re
sponse data were analysis of variance followed by mul
tiple comparison of treatment means and, more recently,
the broken line analysis method which was used in
this study. These procedures assume a linear relation
between weight gain and dietary level of amino acid at
or below the requirement, and when the requirement
is met, the weight gains abruptly plateau and eventu
ally decline if the dietary concentration of the nutrient
exceeds the animals' tolerance for the amino acid. Since
animal responses to increasing dietary concentrations
of essential nutrients do not normally follow such a
pattern, a quadratic, or second-order polynomial regres
sion-analysis method as proposed by Zeitoun, Ulbrey
and Magee (46) would give a more precise empirical fit
of the growth response data. This method will allow
derivation of dietary nutrient concentrations for the
oretical maximum growth and also for a level of re
sponse that is below maximum but within a certain
confidence range, say 95% of maximum. The procedure
of Zeitoun, Ulbrey and Magee (46) would be the pre
ferred method of analysis of growth response data for
practical feeding recommendations.
LITERATURECITED
1. HALVER,I. E., DELONG,D. C. & MERTZ,E. T. (1957) Nutrition
of salmonoid fishes. V. Classification of essential amino acids
for chinook salmon. /. Nutr. 63: 95-105.
2. HALVER,I. E. & SHANKS,W. E. (1960) Nutrition of salmonoid
fishes. VIII. Indispensable amino acids for sockeye salmon. /.
Nutr. 72: 340-346.
3. SHANKS,W. E., GAHIMER,G. D. & HALVER,J. E. (1962) The
indispensable amino acids for rainbow trout. Prog. Fish-Cult. 24:
68-73.
4. DUPREE,H. K. & HALVER,J. E. (1970) Amino acids essential
for the growth of channel catfish, Ictalurus punctatus. Trans.
Am. Fish. Soc. 99: 90-92.
 1546 SANTIAGO AND LOVELL
5. NOSE,T., ARAI,S., LEE,D. & HASHIMOTO,Y. (1974) A note on
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6. YONE,Y. (1976) Nutritional studies of red sea bream. In: Pro
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7. ARAI, S., NOSE,T. & HASHIMOTO,Y. (1972] Amino acids es
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of Warmwater Fishes, 98 pp., National Academy Press, Wash
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9. MAZID, M. A, TANAKA,Y., KATAYAMA,T., SIMPSON,K. L. &
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