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Department of Wildlife and Fisheries Sciences, and Faculty of Nutrition, Texas A&M University System, College Station, TX 77843-2258
3To whom correspondence and reprint requests should be addressed.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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KEY WORDS: • arginine requirement • amino acids • red drum
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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), although there is limited information on
its amino acid requirements. At this time, only the requirements for
lysine (Craig and Gatlin 1992
), total sulfur amino acids
(Moon and Gatlin 1991
) and threonine (Boren and Gatlin 1995
) have been determined for red drum. The dietary
arginine requirement of red drum has not been established, but studies
with other fish species have indicated that arginine is an
indispensable amino acid, and dietary requirements may vary
considerably (NRC 1993
).
The quantitative requirement of some other fish species for arginine
has been determined using diets containing graded levels of arginine,
but all diets were maintained isonitrogenous by adjusting the levels of
aspartic acid and glutamate. Glutamate and glutamine have proven to be
important sources for endogenous arginine in many mammalian species
(Baker 1994
, Blachier et al. 1993
,
Dhanakoti et al. 1990
, Wu and Knabe 1995
). In addition, some fish species (Chiu et al. 1986
) appear to synthesize arginine from dietary glutamate.
Therefore, the objective of this study was to determine the dietary
arginine requirement of juvenile red drum using a mixture of glycine
and aspartate to maintain all diets isonitrogenous, thus limiting the
possibility of dietary glutamate and/or glutamine serving as precursors
for arginine biosynthesis.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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The basal4
and experimental diets contained red drum muscle and crystalline amino
acids to provide 35 g crude protein/100 g diet (Serrano et al. 1992
). Dietary lipid and dextrin were adjusted to provide
13.4 kJ of estimated digestible energy/g diet. The contribution of
arginine to the basal diet by red drum muscle was determined to be 0.65
g/100 g dry diet. In conjunction with the amino acids provided by red
drum muscle, the composition of the amino acid premix was adjusted to
provide amino acid concentrations similar to those provided by 35 g protein/100 g from whole chicken egg or red drum muscle (highest
value), except for arginine. The experimental diets were supplemented
with L-arginine hydrochloride to provide arginine in
increments of 0.30 g/100 g of diet at the expense of a 50:50 mixture
(weight basis) of aspartate and glycine to maintain all diets
isonitrogenous. Dietary arginine concentrations were formulated to
range from 0.65 to 2.75 g/100 g of diet. Amino acid levels in the diets
(except tryptophan, proline and cystine) were determined in duplicate
by reversed-phase HPLC as described by Wu et al. (1997)
. Analyzed arginine levels were within 8% of formulated
values. To minimize the possible effect of variable dietary glutamate
and glutamine, both amino acids were provided solely by red drum muscle
in the diets. Levels of these two amino acids were constant across
diets (totaling 2.5 g/100 g) as confirmed by HPLC analysis. Glycine
levels ranged from 3.4 to 7.5 g/100 g diet. Any differential effects of
glycine as a palatability enhancer (Hughes 1993
) should
have been negligible, especially considering the excellent acceptance
of this type of diet by red drum (Moon and Gatlin 1991
).
The levels of all remaining nutrients were kept constant and met all
established requirements of red drum.
Diets were adjusted to a pH of 7.0 with 6 mol/L NaOH, mechanically
mixed and pressure pelleted as previously described (McGoogan and Gatlin 1998
). The diets were then air-dried and stored
at -20°C (~ 6 wk) until needed. Before being fed, diets were
thawed, crumbled to an appropriate size for ingestion and stored at
4°C (~1 wk).
The feeding trial was conducted in 38-L aquaria configured as closed
recirculating systems with mechanical and biological filtration. The
salinity was maintained at 5 g/L using a synthetic sea salt mixture and
fresh well water. Water temperature was monitored biweekly and averaged
29.1 ± 0.1°C. Aeration was provided to the aquaria via low
pressure electrical blowers and distributed through air stones. Water
quality was monitored weekly for salinity, ammonia, nitrate, nitrite,
hardness and pH, and maintained at optimum levels for red drum
(Neill 1990
) throughout the feeding trial. A diurnal
12-h light:dark cycle was artificially provided by fluorescent lights
controlled by electrical timers.
Groups of juvenile red drum (n = 12) were stocked into the aquaria at an average initial weight of 56.6 ± 0.7 g/group (mean ± SD). These fish had previously been fed the basal diet for 1 wk during which time they became acclimated to the culture conditions and the semipurified diet. Each diet was fed to triplicate groups of fish in a preweighed amount twice daily at a rate that approached apparent satiation. The amount of diet given was calculated on the basis of a percentage of total fish weight per aquarium, and these weights were obtained weekly. The daily rations were reduced gradually by equal amounts among all treatments from 6.5% of total fish weight per day to 5% as the fish grew and their metabolic needs per unit of body weight decreased. The feeding trial was continued for 7 wk. Procedures used in this study were approved by the Texas A&M University System Animal Care and Use Committee.
At the conclusion of the feeding trial, three fish were collected from
each of three replicate aquaria per diet ~15 h after the last
feeding. This postprandial time point was selected on the basis of the
time course of plasma amino acid concentrations of fish fed
casein-caseinate and crystalline amino acid mixes
(Schuhmacher et al. 1995
). Blood was collected from each
fish using heparinized needles and centrifuged (2000 x g, 10 min) for plasma separation. Plasma from three fish
per aquarium was pooled and analyzed for amino acids using HPLC
(Wu and Knabe 1995
). The sampled fish were then killed
with an overdose of 3-aminobenzoic acid ethyl ester (Sigma Chemical,
St. Louis, MO) and frozen (-80°C) before determining carcass
proximate composition. Composite samples of three fish per aquarium at
the beginning and end of the experiment were homogenized and subjected
to proximate analysis as previously described (Serrano et al. 1992
). Protein retention
(PR)5
was determined on the basis of protein gain in fish carcass and dietary
protein consumption.
Weight gain (WG), feed efficiency (FE), protein efficiency ratio (PER),
PR and survival, as well as plasma arginine, glutamate, glutamine,
glycine, citrulline and ornithine concentrations were evaluated as
response criteria and subjected to one-way ANOVA with significance
set at P 
0.05. When significant differences were
detected, means were separated by Duncan’s multiple range test.
Quantitative arginine requirement estimates also were determined by
least-squares regression using the broken-line method
(Robbins 1986
) with respect to the response criteria.
Statistical analyses were performed using the Statistical Analysis
System (SAS 1988
). Values are presented as means (±
SEM).
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RESULTS AND DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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). Weight gain of fish reached a plateau at ~1.85 g arginine/100 g of
diet. Regression analysis of WG data using the broken-line method
estimated an arginine requirement of 1.75 (± 0.18) g/100 g of diet or
5.0 (± 0.5) g/100 g dietary protein. Feed efficiency and PER values
increased with dietary arginine from 0.65 to 1.25 g/100 g diet and then
remained relatively constant at higher levels of arginine intake.
Regression analysis of FE and PER data provided arginine requirement
estimates of 1.44 (± 0.15) and 1.48 (± 0.12) g/100 g diet or 4.1 (±
0.4) and 4.2 (± 0.3) g/100 g dietary protein, respectively. Arginine
did not affect PR (Table 1)
.
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). The differences among plasma arginine levels were significant as they
increased with dietary arginine. In a parallel experiment in this
laboratory (data not shown), juvenile red drum tended (P
= 0.06) to have greater WG and FE values when fed diets
containing suboptimal arginine levels in which glutamate, as opposed to
glycine, was used to maintain diets isonitrogenous. However, data on
circulating glutamate, glutamine, citrulline and ornithine, all of
which are involved in pathways for endogenous synthesis of arginine as
established in other species (Baker 1994
,
Blachier et al. 1993
, Dhanakoti et al. 1990
, Wu and Knabe 1995
), did not provide
evidence to support the occurrence of de novo synthesis of arginine
from either glutamine or glutamate. Further research is required to
determine conclusively whether glutamate may support a compensatory
mechanism for endogenous arginine synthesis during limited dietary
arginine intake in red drum.
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), 3.9 for sea bass
(Dicentrarchus labrax) (Tibaldi et al. 1994
),
4.3 for channel catfish (Ictalurus punctatus)
(Robinson et al. 1981
), 4.3 for common carp
(Cyprinus carpio) (Nose 1979
), 4.4 for hybrid
striped bass (Morone saxatilus x M.
chrysops) (Griffin et al. 1994
), 4.3 for yellow
perch (Perca flavescens) (Twibell and Brown 1997
) and 4.2 for rainbow trout (Onchorynchus
mykiss) (Cho et al. 1992
). However, the arginine
requirement estimate of 5.0 g/100 g dietary protein based on WG in this
study is slightly above the previously reported values but is similar
to the value of 5.3 g/100 g dietary protein reported for milkfish
(Chanos chanos) (Borlongan 1991
). Yellow
perch also showed a higher arginine requirement on the basis of WG
compared with the requirement obtained from FE (Twibell and Brown 1997
); this higher requirement (4.9 g/100 g dietary
protein) is similar to the requirement estimate for red drum based on
WG in this study. On the basis of the various results of this study,
diets for red drum should be formulated to contain arginine at no <4.1
g/100 g dietary protein to ensure rapid growth and efficient feed
utilization.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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2 Funding for J.A.B. was provided in part by the Consejo Nacional para la Ciencia y la Tecnologia (CONACYT-México).
4 Composition of the basal diet (g/100 g dry
weight): 13.2 g red drum muscle (obtained from adult wild fish,
containing 78.0% crude protein and 16.4% lipid, 90.6% dry matter);
19.2 g amino acid premix, dry matter 100.0% [consisting of
L-amino acids from Nutri-Quest, Chesterfield, MO and
providing (g/100 g diet): glycine, 1.49; histidine, 0.69; isoleucine,
1.69; leucine, 2.26; lysine·HCl, 0.69; methionine, 0.89; cystine,
0.71; phenylalanine, 1.55; tyrosine, 1.10; serine, 2.25; threonine,
0.31; tryptophan, 0.42; valine, 1.98; proline, 1.58; alanine, 1.58];
33.4 g dextrin, 94.2% dry matter (United States Biochemical,
Cleveland, OH); 10.0 g menhaden oil, 100.0% dry matter (Omega
Protein, Reedville, VA); 4.0 g mineral premix, 100.0% dry matter
[see McGoogan and Gatlin (1998)]; 3.0 g vitamin premix,
100.0% dry matter [see McGoogan and Gatlin (1998)]; 2.0 g carboxymethyl cellulose, 94.1% dry matter (United States Biochemical); 1.0 g Ca(HPO4)2, 100.0% dry
matter (Fisher Scientific, Pittsburgh); 8.2 g cellulose, 96.4% dry matter (United States Biochemical); 3 g
L-aspartate; 3 g glycine, 100.0% dry matter (Nutri-Quest).
5 Abbreviations used: FE, feed efficiency; PER, protein efficiency ratio; PR, protein retention; WG, weight gain.
Manuscript received August 23, 1999. Revision accepted March 21, 2000.
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REFERENCES |
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