Amino Acids

DOI 10.1007/s00726-008-0171-1

REVIEW ARTICLE

New developments in fish amino acid nutrition: towards functional

and environmentally oriented aquafeeds
Peng Li Æ Kangsen Mai Æ Jesse Trushenski Æ
Guoyao Wu

Received: 25 June 2008 / Accepted: 3 August 2008

Ó Springer-Verlag 2008

Abstract Recent evidence shows that some amino acids balanced aquafeeds to enhance the efficiency and profit-
and their metabolites are important regulators of key ability of global aquaculture production.
metabolic pathways that are necessary for maintenance,
growth, feed intake, nutrient utilization, immunity, Keywords Amino acids
Fish
Health
Growth

behavior, larval metamorphosis, reproduction, as well as Aquafeeds
Aquaculture
resistance to environmental stressors and pathogenic
organisms in various fishes. Therefore, conventional defi- Abbreviations
nitions on essential and nonessential amino acids for fish AA Amino acids
are challenged by numerous discoveries that taurine, glu- GABA c-Aminobutyrate
tamine, glycine, proline and hydroxyproline promote BCAA Branched-chain amino acids
growth, development, and health of aquatic animals. On the HMB Hydroxyl-b-methyl-butyrate
basis of their crucial roles in cell metabolism and physi- NAC N-Acetyl-cysteine
ology, we anticipate that dietary supplementation with NO Nitric oxide
specific amino acids may be beneficial for: (1) increasing NOS Nitric oxide synthase
the chemo-attractive property and nutritional value of P5C Pyrroline-5-carboxylate
aquafeeds with low fishmeal inclusion; (2) optimizing T3 Triiodothyronine
efficiency of metabolic transformation in juvenile and sub- T4 Thyroxine
adult fishes; (3) surpressing aggressive behaviors and
cannibalism; (4) increasing larval performance and sur-
vival; (5) mediating timing and efficiency of spawning;
(6) improving fillet taste and texture; and (7) enhancing Introduction
immunity and tolerance to environmental stresses. Func-
tional amino acids hold great promise for development of Amino acids (AA) are building blocks for protein. On the
basis of needs from diet for growth, AA were traditionally
classified as nutritionally essential (indispensable) or non-
P. Li (&)
G. Wu essential (indispensable) for fish (Table 1). Essential AA
Department of Animal Science, Texas A&M University,
are those that either cannot be synthesized or are inade-
College Station, TX 77843-2471, USA
e-mail: [email protected] quately synthesized de novo by animals relative to needs.
Conditionally essential AA must be provided from the diet
K. Mai under conditions where rates of utilization are greater than
The Key Laboratory of Mariculture (Ministry of Education),
Ocean University of China, 5 Yushan Rd, Qingdao 266003,
rates of synthesis. By definition, all nonessential AA can be
People’s Republic of China synthesized adequately by aquatic animals.
Dietary protein is the major and most expensive com-
J. Trushenski ponent of formulated aquafeeds (Wilson et al. 2002).
Fisheries and Aquaculture Center, Southern Illinois University,
Carbondale, IL 62901-6501, USA
Fishmeal is generally considered to be the most ideal

123
 P. Li et al.

Table 1 Nutritionally essential and nonessential amino acids for fish

and other aquatic animals
Essential AA Nonessential AA Conditionally essential AA

Arginine Alanine Cysteine

Histidine Asparagine Glutamine
Isoleucine Aspartate Hydroxyproline
Leucine Glutamate Proline
Lysine Glycine Taurine
Methionine Serine
Phenylalanine Tyrosine
Threonine
Tryptophan
Valine

protein source for aquatic animals, despite its static global

Fig. 1 Roles of amino acids in growth, development and health of
production, seasonal/geographical variability in quality and fish
composition, and concern as a vector of contamination
(Trushenski et al. 2006). Over the past decades, there have
been rapid increases in global aquaculture and demand, as industry and consumers. Amino acids are potentially
well as price and competition for this valuable feedstuff. important ingredients for these two types of feeds. This
With a growing interest in non-fishmeal protein sources for review summarizes current knowledge on roles of AA in
aquafeeds, it is important to sustain comparable levels of fish nutrition and proposes new strategies to guide devel-
feed intake, feed conversion efficiency, as well as growth opment of AA-balanced aquafeeds.
rate and survival in fishes. Therefore, establishment of
optimal dietary requirements of AA and characterization Alanine, b-alanine, aspartate and asparagine
of alternative protein/AA sources have been a major focus
of fish nutrition research. Alanine and aspartate are the major glucogenic precursors
Emerging evidence from studies with both aquatic and and important energy substrates for fish. In addition,
terrestrial animals shows that many AA regulate key met- aspartate is essential to purine nucleotide synthesis in all
abolic pathways that are crucial to maintenance, growth, cell types. Moreover, alanine is a preferred carrier of
reproduction, and immune responses (Fig. 1). They are nitrogen for inter-organ AA metabolism in fish (Mommsen
termed ‘‘functional AA’’. Identification and dietary sup- et al. 1980). Aspartate and asparagine represent *10% of
plementation of those AA or their biologically active AA in plant and animal proteins, but dietary aspartate may
metabolites is expected to offset adverse effects of be extensively catabolized by the small intestine of fish, as
replacement of fishmeal from aquafeeds, therefore restor- established for mammals. Thus, most of the circulating
ing food intake and growth. Of particular interest, the aspartate is derived from synthesis by skeletal muscle (Wu
‘‘least-cost’’ principle of aquafeed formulation has been 1998). It is noteworthy that aspartate aminotransferase is
evolving in recent years towards customer-based aqua- up-regulated in gill, white and red muscles (Bystriansky
feeds, which has led to the concepts of ‘‘functional et al. 2007), and liver (Aas-Hansen et al. 2005) during
aquafeeds’’ and ‘‘environmentally oriented aquafeeds’’. salinity acclimation or migration.
Functional aquafeeds are defined as feeds supplemented Alanine can stimulate feeding response of certain fishes
with specific ingredient(s) to achieve desirable efficiency although the underlying mechanisms are unknown
of metabolic transformation, growth performance, health, (Shamushaki et al. 2007). Interestingly, dietary supple-
and/or compositional traits of aquacultured animals at mentation with 5% pyruvate can reduce nitrogen excretion
various developmental stages. Environmentally oriented into the surrounding water via enhancement of alanine
aquafeeds are defined as feeds modified to minimize neg- formation, without a negative effect on the growth and
ative impacts of environmental changes (including salinity, vitellogenin synthesis in Atlantic salmon (Olin et al. 1992).
ammonia, extreme temperatures, and stressors imposed by At present, there is a little information regarding supple-
husbandry handling) on growth, health, and reproduction of mentation of aspartate or asparagine to fish diets. Due to
aquacultured animals. Both functional and environmentally rapid oxidation and a lack of toxicity, alanine and aspartate
oriented aquafeeds are designed to meet needs of the are often used to balance nitrogen in fish nutrition research.

123
New developments in fish amino acid nutrition

b-Alanine (a product of aspartate decarboxylation) is proline or polyamines. A substantial decrease in plasma

a constituent of carnosine (b-alanyl-L-histidine), anserine concentrations of arginine and ornithine has been observed
(b-alanyl-1-methyl-L-histidine), carcinine (b-alanyl-hista- for Senegalese sole subjected to acute stress (Aragão et al.
mine), and balenine (b-alanyl-3-methyl-L-histidine). 2008).
Carnosine and anserine are antioxidants and important Except for endothelial NOS (eNOS), genes for inducible
buffers in skeletal muscle of aquatic animals, especially NOS (iNOS) and neuronal NOS (nNOS) have been cloned
migratory pelagic marine fishes (Snyder et al. 2008). Ele- for various fish (Bordieri et al. 2005). While physiological
vated levels of b-alanine in plasma or tissue in response to levels of NO play an important role in cellular signaling, a
acute handling stress or an environmental change have high concentration of NO synthesized by iNOS can result
been reported for fish (Aragão et al. 2008) and oysters in AA and protein modification, oxidative stress, apoptosis,
(Powell et al. 1982); however, the physiological signifi- and killing of target cells (Galli 2007; Mannick 2007).
cance of this finding has not been elucidated. Ogata (2002) Inducible NO production in macrophages is affected by
found that dietary supplementation of b-alanine increased extracellular arginine availability or nutrition of channel
intramuscular concentration of carnosine, but not anserine, catfish (Buentello and Gatlin 1999). nNOS is expressed in
in yellowtail. Kim et al. (2003) observed no positive effect the brain and gill of fish that regulates neurological func-
of b-alanine supplementation on growth performance of tion and development, vascular tone and osmotic pressure
Japanese flounders. Carnosine is abundant in fish but (Hyndman et al. 2006). In addition, arginine can stimulate
absent from plants. Supplementing carnosine to all-plant the release of various hormones such as insulin, growth
protein-based diets did not appear to enhance growth or hormone, and glucagon. Indeed, arginine is a more potent
feed efficiency in rainbow trout (Snyder et al. 2008); activator of insulin release than glucose in fish (Mommsen
however, potential use of carnosine for fish under stressful et al. 2001). Growth- and health-promoting effect of die-
conditions remains to be determined. tary arginine beyond meeting requirement for protein
synthesis has been reported for some fishes. For example,
Arginine, citrulline and ornithine Buentello and Gatlin (2001) found that the survival of
channel catfish in response to challenge with Edwardsiella
Fish have particularly high requirements for dietary argi- ictaluri critically depended upon dietary arginine levels.
nine, because (1) it is abundant in protein (as a peptide- Despite extensive work with mammalian species (Wu and
bound AA) and tissue fluid (as phosphoarginine, a major Morris 1998), little is known about the efficacy of citrulline
reservoir of ATP); and (2) its de novo synthesis is limited to replace arginine in fish diets.
or even completely absent. Elasmobranchs and ureogenic
teleosts may be able to convert citrulline into arginine via Branched-chain AA (BCAA)
argininosuccinate synthase and lyase in liver (Mommsen
et al. 2001). However, it is unknown whether there is net Leucine, isoleucine plus valine account for 18–20% AA in
synthesis of citrulline or arginine by this organ in aquatic plant and animal proteins. As an activator of the target of
animals. There is circumstantial evidence for an arginine- rapamycin (a protein kinase), leucine is considered as a
sparing effect of dietary glutamate in channel catfish functional AA to stimulate muscle protein synthesis and
(Buentello and Gatlin 2000), but this effect may result from inhibit proteolysis in mammals (Nakashima et al. 2007).
an inhibition of arginine degradation. At present, pathways However, such a role for leucine has not been investigated
for arginine synthesis from glutamine, glutamate, proline, for aquatic animals. Choo et al. (1991) reported that
or AA other than citrulline have not been established for excessive leucine increased feed efficiency but depressed
fishes. food intake, growth, and protein deposition of rainbow trout
Arginine is one of the most versatile AA by serving as likely due to antagonism among AA (particularly BCAA).
the precursor for the synthesis of protein, nitric oxide (NO), Notably, in recent years, there has been growing interest in
urea, polyamines, proline, glutamate, creatine and agma- the stimulatory effect of supplemental b-hydroxyl-b-
tine in terrestrial animals (Wu and Morris 1998). Arginine methyl-butyrate (HMB; a leucine metabolite) on growth
also plays a crucial role in regulating endocrine and and immunity in several coldwater aquacultured fishes
reproductive functions, as well as extra-endocrine signaling (Siwicki et al. 2003). However, such a benefit has not been
pathways (including AMP-activated protein kinase and the observed in warm water species, including tilapia and
target of rapamycin) (Jobgen et al. 2006; Yao et al. 2008). hybrid striped bass (Li and Gatlin 2007). This discrepancy
Fish can produce NO by tetrahydrobiopterin-dependent NO may be explained by species differences in leucine metab-
synthase (NOS; Buentello and Gatlin 1999), as well as urea olism among fishes. In addition, studies with fishes (e.g.,
plus ornithine by arginase (Gouillou-Coustans et al. 2002). rainbow trout, Arctic charr, and even shrimp) have revealed
The resultant ornithine can be used for the synthesis of that BCAA can be rapidly mobilized from white muscle and

123
 P. Li et al.

red muscle during exhaustive exercise (Milligan 1997) or feed-grade glutamine and glutamate (e.g., AminoGutTM,
sea water acclimation (Bystriansky et al. 2007), but the rates Ajinomoto Inc., http://www.Ajinomoto.com), it will be of
of leucine release vary greatly among species. practical importance to determine whether this new product
can enhance growth performance and immunity in fish
Glutamate, glutamine, and c-aminobutyrate (GABA) (including shrimp).

Glutamine is one of the most abundant free a-AA in fish Glycine and serine
plasma and muscle, whereas glutamate and its decarbox-
ylation product (GABA) are neurotransmitters present at Glycine and serine are interconverted primarily in liver and
high concentrations in the brain. In addition, GABA is an kidneys by tetrahydrofolate-dependent hydroxymethyl-
inducer of metamorphosis in abalone (Morse et al. 1979). transferase. These two AA participate in gluconeogenesis,
Moreover, glutamine is essential for the synthesis of purine sulfur AA metabolism, one-carbon unit metabolism, and
and pyrimidine nucleotides in all cells. Through renal fat digestion (Fang et al. 2002), and also stimulate feed
ammoniagenesis, glutamine also plays an important role in intake by many fishes (Shamushaki et al. 2007). Glycine
regulating the acid–base balance in the body. Further, may function to regulate gene expression in fish, as dietary
glutamine stimulate muscle protein synthesis in mammals supplementation with glycine has been shown to increase
(Wu et al. 2007), but such information is not yet available hepatic thyroxine 50 monodeiodinase activity in rainbow
for fish. As a major energy substrate for leukocytes and a trout (Riley et al. 1996), therefore enhancing the efficiency
key modulator of cytokine and NO production, glutamine of nutrient absorption and anabolic events. Interestingly,
is crucial to the immune response in fish (Buentello and glycine has a critical role in the osmoregulatory responses
Gatlin 1999; Li et al. 2007). of fishes and shellfishes (e.g., oysters) to environmental
Glutamate and glutamine represent *20% of AA in plant stress. For example, oysters rapidly take up free glycine
and animal proteins, but they could be completely degraded from the surrounding water, and they synthesize gill gly-
by the gut of aquatic animals, as reported for terrestrial cine in response to rapid changes in salinity or anoxia
mammals (Wu 1998). Thus, most of glutamate and gluta- (Powell et al. 1982). Importantly, glycine-enrichment can
mine in plasma may be synthesized from BCAA and a- enhance the survival of oysters after being transferred to
ketoglutarate in skeletal muscle. Glutamate is a substrate for freshwater from seawater (Takeuchi 2007) and may be an
glutamine synthesis by ATP-dependent glutamine synthe- innovative method to improve oyster larviculture. Simi-
tase, whereas glutamine is hydrolyzed by phosphate- larly, tissue levels of serine in aquatic animals (including
dependent glutaminase to generate glutamate (Anderson shrimp) are increased by abrupt salinity change (Silvia
et al. 2002). This intracellular and intercellular glutamine– et al. 2002) or hardness change (Buentello and Gatlin
glutamate cycle is well known for mammals, but has not 2002). At present, little is known about whether dietary
been firmly established for fishes. As major enzymes initi- serine supplementation may optimize fish survival and
ating glutamate degradation, glutamate dehydrogenase is growth in suboptimal culture conditions.
expressed in fish liver and white muscle, whereas glutamate
transaminase is abundant in all fish tissues (Wilson 2002). Histidine
Glutamine synthetase is abundant in brain, intestine, liver,
red muscle, gill/kidney, white muscle, and heart of fish, and Histidine is an abundant AA in plasma albumin of fish
the hepatic protein is up-regulated by cortisol (Vijayan et al. (Szebedinszky and Gilmour 2002). It is also rich in fish
1996) or high environmental ammonia (Anderson et al. muscle as a free AA or carnosine. Histidine participates in
2002). Both glutamate and glutamine are important energy one-carbon unit metabolism, therefore affecting DNA and
substrates in fish, but tissue-specific metabolism of the two protein synthesis. In addition to serving as an energy fuel
AA is fully defined for aquatic animals. during starvation, histidine is a major component of non-
High dietary levels of GABA inhibits food intake by carbonate buffers protecting fish against changes in pH
Japanese flounder (Kim et al. 2003). In contrast, studies with resulting from hypoxia, burst-swimming, and increased
Atlantic salmon indicate that dietary supplementation with lactacidosis. The capacity of the non-carbonate buffer sys-
5% a-ketoglutarate can reduce nitrogen excretion into the tem among various fish species varies considerably, which
water without a negative effect on growth and vitellogenin is likely attributable to long-term adaptation to environ-
synthesis (Olin et al. 1992). Importantly, dietary glutamine ments. Interestingly, intramuscular histidine concentration
supplementation enhances weight gain, feed intake, gain:- is substantially increased before spawning migration of
feed ratio, intestinal development, and digestive enzyme salmon (Mommsen et al. 1980). It is possible that histidine
activities in Jian carp (Lin and Zhou 2006). With the metabolism and its dietary requirement in fish are regulated
recent commercial availability of commercial mixtures of by many environmental and endocrine factors.

123
New developments in fish amino acid nutrition

Because histidine and its related imidazole derivatives Phenylalanine and tyrosine
(anserine and carnosine) confer desirable taste and texture
(e.g., sweetness, heaviness and thickness), dietary supple- Phenylalanine can be converted to tyrosine by tetrahydro-
mentation of histidine can improve sensory attributes (e.g., biopterin-dependent phenylalanine hydroxylase in liver and
flavor) of aquacultured seafoods (Ogata 2002). Addition- kidneys. Thus, adding tyrosine to diets for fish can reduce
ally, Førde-Skjærvik et al. (2006) reported that dietary requirement for phenylalanine. Research on supplementing
supplementation of histidine increased intramuscular his- phenylalanine and tyrosine to aquafeeds and their potential
tidine levels and pH, while reducing muscle gapping in influences on aquatic animals is currently limited. Tyrosine
Atlantic cod post-mortem. This, in turn, contributes to a is a common precursor for important hormones and neu-
higher quality fillet. rotransmitters, including thyroxine (T4), triiodothyronine,
epinephrine, norepinephrine, dopamine, and melanin. These
Lysine, carnitine and cadaverine molecules have important regulatory roles (Chang et al.
2007; Yoo et al. 2000); hence, dietary levels of phenylala-
Lysine is often one of the most limiting AA in ingredi- nine and tyrosine could profoundly influence pigmentation
ents used for production of commercial fish feeds, development, feed intake, growth performance, immunity,
especially when fishmeal is replaced by plant protein and survival of fish in natural environment. Consequently,
sources (Mai et al. 2006a). Therefore, dietary lysine dietary requirements for phenylalanine and tyrosine of fish
levels critically affect fish growth performance and increase substantially during metamorphism (Pinto et al.
health. With the commercial availability of feed-grade 2008). Additionally, oral administration of T4 to carp,
lysine, its addition to plant protein-based diets allows for channel catfish and flounder fish increased protein digest-
cost-effective reduction of dietary crude protein without ibility, digestive enzyme activity, nutrient retention, growth
affecting fish growth performance (Mai et al. 2006a). rate, and feed efficiency (Garg 2007).
This nutritional strategy can also decrease the excretion
of ammonia and soluble phosphorus from fish (Cheng Polyamines
et al. 2003). In addition, dietary lysine supplementation
is effective in enhancement of immune responses and Polyamines (putrescine, spermidine, and spermine) are
gastrointestinal development of agastric fish (Jian carp) naturally occurring polycationic substances that are
(Zhou 2005). essential for cell proliferation and differentiation. They are
Lysine is a substrate for the synthesis of carnitine, which synthesized from arginine- or proline-derived ornithine in
is required for the transport of long chain fatty acids from mammals (Wu et al. 2008); however, these pathways have
the cytosol into mitochondria for oxidation. Potential not been established for fish. Spermine is relatively abun-
benefits of dietary carnitine supplementation include dant in most fishes, possibly indicating active polyamine
growth promotion, protection against the toxicity of synthesis or limited degradation. Natural dietary sources of
ammonia and xenobiotics, improved acclimation to polyamines for fishes have not been fully defined. It has
extreme temperature changes and associated stress, and been postulated that gastrointestinal microbes in fish larvae
enhanced reproduction performance (Harpaz 2005). How- can produce polyamines and facilitate development of
ever, because of differences in experimental conditions, the small intestine. Early study indicated that dietary
data on the nutritional value and potential economical supplementation of putrescine did not enhance growth
return of carnitine supplementation have not been consis- performance of juvenile rainbow trout (Cowey and Cho
tent for various types of fishes and even within the same 1992). However, supplementing spermine to micropartic-
species. Further research is necessary to explain this dis- ulate diets can increase digestive enzyme activity and
crepancy and identify an effective dose of carnitine and promote intestinal maturation, therefore enhancing the
responsive conditions. survival of larval fish (e.g., European sea bass) (Péres et al.
Cadaverine is a decarboxylation product of lysine during 1997). It should be noted that high doses of polyamines are
putrefaction of animal tissue, such as fishmeal staling. toxic to aquatic animals and reduce growth performance
Tapia-Salazar et al. (2004) reported that combined sup- (Cowey and Cho 1992).
plementation of cadaverine and histamine can enhance feed
consumption and growth of blue shrimp, rather than being Proline and hydroxyproline
responsible for the reduced nutritional value of stale fish-
meal as suggested by Cowey and Cho (1992). Histamine Proline is traditionally thought to be a dispensable AA for
can regulate blood flow and intestinal development in fish and promotes feed intake. In mammals, proline is
mammals (Ou et al. 2007), but these effects have not been synthesized from arginine, ornithine, glutamine, and glu-
investigated for aquatic animals. tamate in a tissue-specific manner (Wu and Morris 1998).

123
 P. Li et al.

However, pathways for proline synthesis in fish are not taurine (Gaylord et al. 2007). Transcription of other
well defined. Indeed, whether proline can be sufficiently enzyme, cysteine sulfinate decarboxylase, in the liver of
synthesized in fish to meet requirements for maintenance, rainbow trout is not influenced by dietary taurine, but
maximal growth or optimal health remains largely inhibited by dietary methionine (Gaylord et al. 2007).
unknown. Dabrowski et al. (2005) reported the presence of It would be important to determine whether changes
pyrroline-5-carboxylate (P5C) reductase in the liver and in maximum activities of enzymes measured at saturated
intestine of rainbow trout, but the source of P5C was not concentrations of substrates translate into alterations in
identified. These authors also found that proline concen- the flux of methionine to taurine. S-Adenosylmethionine,
tration in muscle of rainbow trout alevin was dependent on which is an important methyl group donor for creatine
dietary proline and that endogenous synthesis of proline and spermidine synthesis (Grillo and Colombatto 2007), is
from glutamate could not meet the requirement for proline crucial for metabolic regulation in fish. However, it is not
(Zhang et al. 2006). Therefore, proline is now considered known whether dietary supplementation with S-adenosyl-
as a conditionally essential AA for fish in early life stage methionine may enhance fish survival or growth.
and perhaps also for adult fish. Methionine and its derivatives are produced commer-
Hydroxyproline content in fish meal is much higher than cially by chemical processes. Methionine is commonly
that in plants proteins. In animals, hydroxyproline is pro- available in the DL-form. L-Methionine, the natural isomer,
duced via the post-translational hydroxylation of proline in is absorbed readily and used efficiently by animals.
protein (primarily collagen) by vitamin C-dependent prolyl D-Methionine must be transaminated into a-ketoacid by
hydroxylase. Fish skin and bones are important sources of D-methionine oxidase, and the a-ketoacid is then converted
collagen. Female fish (e.g., ayu) experience considerable into L-methionine by transaminases (Wu and Thompson
collagen demobilization and lose hydroxyproline during 1989). In contrast to mammals, the utilization of methionine
maturation and spawning (Toyohara et al. 1997). hydroxyl analog (a synthetic non-nitrogenous compound)
Hydroxyproline has been demonstrated as the only free AA by fishes or crustaceans is not highly efficient (Wilson
in tissues that is positively correlated to the growth rate of 2002). Of particular interest, because of the presence of a
juvenile salmon (Sunde et al. 2001). Hydroxyproline may high deacetylase activity in the cytosol of animal tissue,
also be a signaling molecule that regulates cellular redox N-acetyl-methionine is an excellent precursor of sulfur AA
state and apoptosis (Phang et al. 2008). Recently, Aksnes in fish (Keembiyehetty and Gatlin 1995).
et al. (2008) found that dietary supplementation of There is limited information about nutritional essenti-
hydroxyproline, but not proline, increased growth rate ality of cysteine for fish. It has been estimated that cysteine
and modified bone composition of salmon. Although the can spare 40–60% of methionine in the diets for various
underlying mechanisms are unknown, these findings sup- fishes (Wilson 2002). Dietary supplementation with N-
port the notion that AA derivatives are potent growth acetyl-cysteine (NAC) is an effective means to augment the
promoters for fish. Addition of hydroxyproline to plant provision of cysteine, therefore enhancing the synthesis
protein-based aquafeeds may be beneficial for maximizing of glutathione (a major antioxidant tripeptide in cells) in
survival and growth of aquatic animals. animals (Wu et al. 2004). Some studies reveal that adding
glutathione to diets can spare 75% requirement of methi-
Sulfur amino acids: methionine, cysteine and taurine onine by hybrid striped bass (Keembiyehetty and Gatlin
1995), suggesting that glutathione synthesis is an important
Methionine is usually the first limiting AA in many fish metabolic fate of methionine. Interestingly, dietary NAC
diets, especially those containing high levels of plant can reduce cutaneous mucus viscosity, however, it failed
protein sources, such as soybean meal, peanut meal, and to confer protection to rainbow trout or Atlantic salmon
corpa meal (Mai et al. 2006b). Pathways of methionine infection by Paramoeba spp. (amoebic gill disease)
transmethylation, remethylation, transsulfuration for the infection (Powell et al. 2007).
synthesis of cysteine and taurine are known for mammalian Taurine is not incorporated into proteins, but plays
liver, and are likely present in fish despite possible quan- important roles in fat digestion, antioxidative defense, cel-
titative differences among species. For example, the lular osmoregulation, as well as development of visual,
conversion of methionine into taurine has been reported neural and muscular systems (Fang et al. 2002; Omura and
for fish (Wilson 2002), but little is known about the Inagaki 2000). Taurine is abundant in fish meal and animal
intermediates, enzyme activities, as well as hormonal and products (particularly marine invertebrate products), but
nutritional regulation. Interestingly, cysteine dioxygenase, absent from plants. Convincing evidence shows that sup-
one of the two rate-controlling enzymes in taurine syn- plementing taurine to all-plant protein diets can promote
thesis from cysteine, is up-regulated by dietary methionine growth and efficiency of feed utilization by carnivore fish,
at the transcriptional level, but not affected by dietary such as rainbow trout (Gaylord et al. 2007) and Japanese

123
New developments in fish amino acid nutrition

flounder (Kim et al. 2003), but not common carp (Kim et al. role in mediating testicular maturation of certain fish,
2008), suggesting suboptimal de novo synthesis of taurine including masu salmon (Amano et al. 2004) and Indian
by certain species. Additionally, taurine concentration in carp (Bhattacharya et al. 2007). Exogenous administration
tissues of certain fish is greatly influenced by changes in of melatonin to fish can increase plasma concentrations and
salinity (Dabrowski et al. 2005) or hardness (Buentello and also stimulate testicular maturation. It should be borne in
Gatlin 2002). However, potential impacts of these envi- mind that the dose, timing, photoperiod and season are
ronmental factors on sulfur AA metabolism, utilization and important factors that affect the efficacy of exogenous
requirement are unknown. Nonetheless, dietary supple- melatonin in mediating fish reproductive performance.
mentation of taurine can promote intestinal development of
cobia larvae, which may provide a new means to improve
efficiency of larviculture (Salze et al. 2008). Challenges of uses of AA in aquafeeds

Threonine There are technical difficulties in the use of crystalline AA

for aquafeeds. A primary concern is leaching of free AA
Most of the previous research on threonine was focused on into water environment, as well as their degradation by
minimum dietary requirements for various fishes, because enterocytes and/or microflora in the gastrointestinal tract
of its deficiency in plant proteins. Dietary threonine levels (Wu 1998). Second, differences in rates of intestinal
can affect immunity in mammals (Li et al. 2007). In absorption between synthetic and protein-bound AA may
addition, threonine is the major component of mucin in reduce the efficiency of crystalline AA (Murai et al. 1982).
the small intestine, therefore regulating intestinal barrier A wide variety of microbinding processes have been
integrity and function. However, potential use of threonine developed to ameliorate these problems. However, most of
in aquafeed beyond meeting the dietary requirement has the available methods were either registered or patented,
received little attention. instead of being published in peer-reviewed journals. In
addition, there may be species-specific response to the
Tryptophan, serotonin and melatonin microencapsulation process. For example, certain binding
with lipids causes low utilization of arginine by abalone,
Tryptophan can be converted to serotonin (5-hydroxy- probably because of low lipase activity in the abalone gut
tryptamine; a neurotransmitter) and melatonin (an and subsequent failure to degrade the fat coating arginine
antioxidant) (Fang et al. 2002). Aggressive interactions and particles (Britz et al. 1997). Third, supplementing AA to
cannibalism of carnivore fish may cause substantial pro- aquafeeds (especially purified diets) may influence acid–
duction losses under conditions of intensive rearing. base and electrolyte balances, ratios of AA in intestinal
Chronic increase in brain concentrations and turnover lumen and plasma, as well as digestion and absorption of
of serotonin are associated with suppressed aggression. nutrients. Fourth, the Maillard reaction in the process of
Dietary supplementation with L-tryptophan can inhibit feed extrusion can substantially reduce the bioavailability
aggression in juvenile rainbow trout (Hseu et al. 2003), of free AA, especially lysine and arginine (Csapó et al.
reduce cannibalism and stress-induced anorexia in juvenile 2008). Fifth, pathways of AA metabolism and its regulation
grouper (Höglund et al. 2007), and prevent a stress-induced by neural, endocrine, and environmental factors in aquatic
cortisol surge (Lepage et al. 2003). Because prolonged animals are poorly understood, therefore limiting devel-
elevation of cortisol negatively affects growth, feed intake, opment of growth-promoting means through targeted
protein accretion, immunity and disease challenge (Vijayan metabolic control. Sixth, research on technologies that
et al. 1996), use of tryptophan may be a promising nutri- allow efficient delivery of limiting AA or functional AA
tional strategy for health management in aquaculture (e.g., to fish larvae through live feeds is still limited (Saavedra
transport, handling, and vaccination). et al. 2008). Overcoming those obstacles would result
Brain serotonin is synthesized in neurons, while in a substantial improvement in utilization efficiency of
peripheral serotonin is formed and released by gut mucous supplemental AA for the formulation of aquafeeds.
in response to different stimuli such as gastric distension,
hyperosmotic solutions or presence of glucose. Both
endogenous and exogenous (dietary) serotonin can inhibit Conclusion and perspectives
food intake by fish (e.g., European sea bass) and terrestrial
animals (Rubio et al. 2006). Melatonin is produced in the Amino acids play important and versatile roles in fish
pineal organ, retina and gastrointestinal tract of fish, and is nutrition and metabolism. These functions include cell sig-
degraded in liver. The synthesis of melatonin in fish pineal naling (e.g., NO, arginine, glutamine, leucine, proline, and
organ is regulated by photoperiod and plays an important polyamines); appetite stimulation (e.g., alanine, glutamate,

123
 P. Li et al.

Table 2 Roles of amino acids in physiological functions and metabolism of aquatic animals
Amino acid Product Function Species Reference

Amino acids Various proteins Structure, transport, regulation, All animals Li et al. (2007)
immunity, signaling, and fuels
Ala, Glu and Ser Directly Appetite Many fishes Shamushaki et al. (2007)
Arg NO Kill invaded microorganisms Channel Catfish Buentello and Gatlin (1999)
Arg NO Facilitate neurological function Tilapia Bordieri et al. (2005)
and development
Arg NO Regulate vascular tone, blood Killifish Hyndman et al. (2006)
flow, osmolarity in gill, and cell
signaling
Arg and Met Spermine Induce larval intestinal maturation Sea bass Péres et al. (1997)
Arg, Met, and Gly Creatine High energy storage; antioxidant Arctic charr Bystriansky et al. (2007)
Cys, Glu, and Gly Glutathione Antioxidant and cell signaling All animals Wu et al. (2004)
Glu and Gln Directly Ammonia removal Rainbow trout Anderson et al. (2002)
Glu GABA Promote metamorphosis Abalone Morse et al. (1979)
Glu GABA Regulate food intake Japanese flounder Kim et al. (2003)
Gln Directly Increase growth, feed efficiency Carp Lin and Zhou (2006)
and gut development
Gln Directly Fuel for macrophage; Cell Channel catfish Buentello and Gatlin (1999)
signaling
Gln, Gly, and Asp Nucleotides Genetic information storage and Various fishes Li and Gatlin (2006)
expression, biosynthesis,
immunity and reproduction
Gly Directly Increase hepatic T4 Rainbow trout Riley et al. (1996)
50 monodeiodinase
Gly Directly Osmoregulation Oyster Takeuchi (2007)
His Directly and carnosine Protection against pH change Salmon Mommsen et al. (1980)
Leu HMB Immunity modulation; Cell Various fishes Li and Gatlin (2007)
signaling
Lys and Met Carnitine Lipid transporter on mitochondrial Various fishes Harpaz (2005)
membrane
Met Choline Structure in membrane; Various fishes Mai et al. (2006b)
neurotransmitter; betaine
synthesis
Proline P5C Redox regulation; Cell signaling Possibly in fish Phang et al. (2008)
Proline Hydroxyproline Enhance growth; Collagen Salmon Aksnes et al. (2008)
function
Phe and Tyr T4, T3 Influence metamorphosis Sole Pinto et al. (2008)
Phe and Tyr T4, T3 Enhance growth performance Channel catfish Garg (2007)
Phe and Tyr T4, T3 Influence pigmentation Japanese flounder Yoo et al. (2000)
Melanin Influence pigmentation Rainbow trout Boonanuntanasarn et al. (2004)
Phe and Tyr Epinephrine, norepinephrine Neurotransmitters that modulate Flounder Damasceno-Oliveira et al. (2007)
stress responses
Phe and Tyr Dopamine Down-regulated immunity Shrimp Chang et al. (2007)
Trp Serotonin Modulate cortisol release, behavior Rainbow trout Lepage et al. (2003)
and feeding
Trp Melatonin Improve testicular development Masu salmon Amano et al. (2004)
Taurine Directly Osmotic pressure regulation Carp Zhang et al. (2006)
Taurine Directly Hardness adaptation Channel catfish Buentello and Gatlin (2002)
Taurine Directly Gut development Cobia Salze et al. (2008)
Taurine Directly Retinal development Glass eel Omura and Inagaki (2000)
HMB hydroxyl-b-methyl-butyrate; NO nitric oxide; P5C pyrroline-5-carboxylate; T3 triiodothyronine; T4 thyroxine

123
New developments in fish amino acid nutrition

proline, and serine); growth and development regulation to a high exogenous ammonia concentration. J Exp Biol
(e.g., arginine, glutamine, hydroxyproline, leucine, and T4); 205:2053–2065
Aragão C, Corte-Real J, Costas B, Dinis MT, Conceição LEC (2008)
energy utilization (e.g., NO, T4, and carnitine); immunity Stress response and changes in amino acid requirements in
(e.g., NO, arginine, glutamine, and dopamine); osmoregu- Senegalese sole (Solea senegalensis Kaup 1858). Amino Acids
lation (e.g., glycine, taurine, b-alanine, and arginine); 34:143–158
ammonia detoxification (e.g., glutamate, glutamine, and Bhattacharya S, Chattoraj A, Maitra SK (2007) Melatonin in the
regulation of annual testicular events in carp catla catla:
citrulline); antioxidative defense (e.g., glutathione, cysteine, evidence from the studies on the effects of exogenous melatonin,
glutamine, glycine, and taurine); metamorphosis (e.g., continuous light, and continuous darkness. Chronobiol Int
tyrosine, T4, and GABA); pigmentation (e.g., T4 and mel- 24:629–650
anin); gut development (e.g., taurine, glutamine, arginine, Boonanuntanasarn S, Yoshizaki G, Iwai K, Takeuchi T (2004)
Molecular cloning, gene expression in albino mutants and gene
threonine, and polyamines); neuronal development (e.g., knockdown studies of tyrosinase mRNA in rainbow trout.
NO, arginine, taurine, and creatine); stress responses (e.g., Pigment Cell Res 17:413–421
tryptophan, serotonin, branched-chain AA and glutamine); Bordieri L, di Patti MCB, Miele R, Cioni C (2005) Partial cloning of
reproduction (e.g., NO, polyamines, arginine, melatonin, neuronal nitric oxide synthase (nNOS) cDNA and regional
distribution of nNOS mRNA in the central nervous system of the
and hydroxyproline); and suppression of aggressive behav- Nile tilapia Oreochromis niloticus. Mol Brain Res 142:123–133
ior (e.g., tryptophan and serotonin) in aquatic animals Britz PJ, Bacela N, Hecht T (1997) Can crystalline arginine be used to
(Table 2). In addition, certain AA (glutamate, histidine, and quantify the arginine requirement of abalone? Aquaculture
glycine) influence taste, texture, and even post-mortem 157:95–105
Buentello JA, Gatlin DM (1999) Nitric oxide production in activated
seafood quality. Dietary supplementation of AA or deriva- macrophages from channel catfish (Ictalurus punctatus): influ-
tives, modulation of their metabolism, or a combination of ence of dietary arginine and culture media. Aquaculture 179:513–
the two may provide new strategies to develop AA-balanced 521
feeds that can offset environmental impacts on aquacultured Buentello JA, Gatlin DM (2000) The dietary arginine requirement of
channel catfish (Ictalurus punctatus) is influenced by endoge-
animals, improve growth performance, and profitability of nous synthesis of arginine from glutamic acid. Aquaculture
the aquaculture industry. Compelling evidence shows that 188:311–321
dietary supplementation of L-lysine HCl, DL-methionine, Buentello JA, Gatlin DM (2001) Effects of elevated dietary arginine
threonine, and tryptophan to compensate for their deficien- on resistance of channel catfish to exposure to Edwardsiella
ictaluri. J Aquat Animal Health 13:194–201
cies in plant feedstuffs, is highly cost-effective for many fish Buentello JA, Gatlin DM (2002) Preliminary observations on the effects
species. Additionally, achieving large-scale feed-grade of water hardness on free taurine and other amino acids in plasma
arginine and glutamine hold great promise for growth and and muscle of channel catfish. North Am J Aquac 64:95–102
health management in aquaculture. The coming decade will Bystriansky JS, Frick NT, Ballantyne JS (2007) Intermediary
metabolism of Arctic char Salvelinus etabol during short-term
witness continuing advances in AA nutrition technologies salinity exposure. J Exp Biol 210:1971–1985
and their application to formulate functional and environ- Csapó J, Varga-Visi E,_ Lóki K, Albert CS, Salamon SZ (2008) The
mentally oriented aquafeeds. influence of extrusion on loss and racemization of amino acids.
Amino Acids 34:287–292
Acknowledgments We thank Drs. Joe Fox and Gibson Gaylord for Chang CC, Wu ZR, Kuo CM, Cheng W (2007) Dopamine depress
helpful comments on this work, as well as Frances Mutscher for office immunity of tiger shrimp Penaeus Monodon. Fish Shellfish
support. Immunol 24:24–33
Cheng ZJ, Hardy RW, Usry JL (2003) Plant protein ingredients with
lysine supplementation reduce dietary protein levels in rainbow
trout Oncorhynchus mykiss diets, and reduce ammonia nitrogen
References and phosphorus excretion. Aquaculture 218:553–565
Choo P, Smith TK, Cho CY, Ferguson HW (1991) Dietary excesses
Aas-Hansen O, Vijayan MM, Johnsen HK, Cameron C, Jorgensen EH of leucine influence growth and body composition of rainbow
(2005) Resmoltification in wild, anadromous Arctic char (Salv- trout. J Nutr 121:1932–1939
elinus etabol): a survey of osmoregulatory, metabolic, and Cowey CB, Cho CY (1992) Failure of dietary putrescine to enhance
endocrine changes preceding annual seaward migration. Can J the growth of rainbow trout (Oncorhynnchus mykiss). Can J Fish
Fish Aquat Sci 62:195–204 Aquat Sci 94:2473–2496
Aksnes A, Mundheim H, Toppe J, Albrektsen S (2008) The effect Dabrowski K, Terjesen BF, Zhang YF, Phang JM, Lee KJ (2005) A
of dietary hydroxyproline supplementation on salmon (Salmo concept of dietary dipeptides: a step to resolve the problem of
salar L.) fed high plant protein diets. Aquaculture 275:242– amino acid availability in the early life of vertebrates. J Exp Biol
249 208:2885–2894
Amano M, Iigo M, Ikuta K, Kitamura S, Okuzawa K, Yamada H, Damasceno-Oliveira A, Fernandez-Duran B, Goncalves J, Serrao P,
Yamamori K (2004) Disturbance of plasma melatonin profile by Soares-da-Silva P, Reis-Henriques MA, Coimbra J (2007)
high dose melatonin administration inhibits testicular maturation Effects of cyclic hydrostatic pressure on the brain biogenic
of precocious male masu salmon. Zool Sci 24:79–85 amines concentrations in the flounder, Platichthys flesus. Gen
Anderson PM, Broderius MA, Fong KC, Tsui KNT, Chew SF, Ip YK Comp Endocrinol 153:385–389
(2002) Glutamine synthetase expression in liver, muscle, stom- Fang YZ, Yang S, Wu G (2002) Free radicals, antioxidants, and
ach and intestine of Bostrichthys sinensis in response to exposure nutrition. Nutrition 8:872–879

123
 P. Li et al.

Førde-Skjærvik O, Skjærvik O, Mørkøre T, Thomassen MS, Rørvik Mai K, Wan J, Ai Q, Xu W, Liufu Z, Zhang L, Zhang C, Li H (2006b)
KA (2006) Dietary influence on quality of farmed Atlantic cod Dietary methionine requirement of juvenile yellow croaker
(Gadus morhua): effect on glycolysis and buffering capacity in Pseudosciaena crocea R. Aquaculture 251:564–572
white muscle. Aquaculture 252:409–420 Mannick JB (2007) Regulation of apoptosis by protein S-nitrosyla-
Garg SK (2007) Effect of oral administration of l–thyroxine (T4) on tion. Amino Acids 32:523–526
growth performance, digestibility, and nutrient retention in Milligan CL (1997) The role of cortisol in amino acid mobilization
Channa punctatus (Bloch) and Heteropneustes fossilis (Bloch). and metabolism following exhaustive exercise in rainbow trout
Fish Biochem Physiol 33:347–358 (Oncorhychus mykiss Walbaum). Fish Physiol Biochem 16:119–
Gaylord TG, Barrows FT, Teague AM, Johansen KA, Overturf KE, 128
Shephed B (2007) Supplementation of taurine and methionine to Mommsen TP, French CJ, Hochachka PW (1980) Sites and patterns
all-plant protein diets for rainbow trout (Oncorhynchus mykiss). of protein and amino acid utilization during the spawning
Aquaculture 269:514–524 migration of salmon. Can J Zool 58:1785–1799
Gouillou-Coustans MF, Fournier V, Métailler R, Vachot C, Des- Mommsen TP, Moon TW, Plisetskaya EM (2001) Effects on arginine
bruyères E, Huelvan C, Moriceau J, Le Delliou H, Kaushik SJ on pancreatic hormones and hepatic metabolism in rainbow
(2002) Dietary arginine degradation is a major pathway in trout. Physiol Biochem Zool 74:668–678
ureagenesis in turbot (Psetta Maxima). Comp Biochem Physiol Morse DE, Hooker N, Duncan H, Jensen L (1979) c-Aminobutyric
A 132:305–319 acid, a neurotransmitter, induces planktonic abalone larvae to
Galli F (2007) Amino acid and protein modification by oxygen and settle and begin metamorphosis. Science 204:407
nitrogen species. Amino Acids 32:497–499 Murai T, Akiyama T, Ogata H, Hirasawa Y, Nose T (1982) Effect
Grillo MA, Colombatto S (2007) S-Adenosylmethionine and radical- of coating amino acids with casein supplemented to gelatin diet
based catalysis. Amino Acids 32:197–202 on plasma free amino acids of carp. Bull Jpn Sot Sci Fish
Harpaz S (2005) L-Carnitine and its attributed functions in fish culture 48:03–710
and nutrition—a review. Aquaculture 249:3–21 Nakashima K, Yakabe Y, Ishida A, Yamazaki M, Abe H (2007)
Höglund E, Sorensen C, Bakke MJ, Nilsson GE, Øverli Ø (2007) Suppression of myofibrillar proteolysis in chick skeletal muscles
Attenuation of stress-induced anorexia in brown trout (Salmo by a-ketoisocaproate. Amino Acids 33:499–503
trutta) by pre-treatment with dietary L-tryptophan. Br J Nutr Ogata HY (2002) Muscle buffering capacity of yellowtail fed diets
97:786–789 supplemented with crystalline histidine. J Fish Biol 61:1504–
Hseu JR, Lu FI, Su HM, Wang LS, Tsai CL, Hwang PP (2003) Effect 1512
of exogenous tryptophan on cannibalism, survival and growth in Olin T, Bergstrom E, Jungvid H, Vonderdecken A (1992) Effect of
juvenile grouper, Epinephelus coioides. Aquaculture 218:251– dietary keto acids on intermediary metabolism of nutrients in
263 Atlantic salmon (Salmo salar) during 17-b estrodial induced
Hyndman KA, Choe KP, Havird JC, Rose RE, Piermarini PM, Evans vitellogenin synthesis. Acta Agric Scandinavica A 42:246–253
DH (2006) Neuronal nitric oxide synthase in the gill of killifish Omura Y, Inagaki M (2000) Immunocytochemical localization of
(Fundulus heteroclitus). Comp Biochem Physiol B 144:510–519 taurine in the fish retina under light and dark adaptations. Amino
Jobgen WS, Fried SK, Fu WJ, Meininger CJ, Wu G (2006) Acids 19:593–604
Regulatory role for the arginine-nitric oxide pathway in metab- Ou DY, Li DF, Cao YH, Li XL, Yin JD, Qiao SY, Wu G (2007)
olism of energy substrates. J Nutr Biochem 17:571–588 Dietary supplementation with zinc oxide decreases expression of
Keembiyehetty CN, Gatlin DM (1995) Evaluation of different sulfur- the stem cell factor in the small intestine of weanling pigs. J Nutr
compounds in the diet of juvenile sunshine bass (Morone Biochem 18:820–826
chrysops 9 M. saxatillis). Comp Biochem Physiol A 112:155– Péres A, Cahu CL, Zambonino-Infante JL (1997) Dietary spermine
159 supplementation induces intestinal maturation in sea bass
Kim SK, Takeuchi T, Yokoyama M, Murata Y (2003) Effect of (Dicentrarchus labrax) larvae. Fish Biochem Physiol 16:479–
dietary supplementation with taurine, b-alanine, and GABA 485
on the growth of juvenile and fingerling Japanese flounder Phang JM, Donald SP, Pandhare J, Liu Y (2008) The metabolism of
Paralichthys olivaceus. Fish Sci 69:242–248 proline, as a stress substrate, modulates carcinogenic pathways.
Kim SK, Matsunari H, Takeuchi T, Yokoyama M, Furuita H, Murata Amino Acids. doi:10.1007/s00726-008-0063-4
Y, Goto T (2008) Comparison of taurine biosynthesis ability Pinto W, Figueira L, Dinis MT, Aragão C (2008) How does fish
between juveniles of Japanese flounder and common carp. metamorphosis affect aromatic amino acid metabolism? Amino
Amino Acids 35:161–168 Acids doi:10.1007/s00726-008-0045-6
Lepage O, Tottmar O, Winberg S (2003) Elevated dietary intake of Powell EN, Kasschau M, Chen E, Koenig M, Pecon J (1982) Changes
L-tryptophan counteracts the stress-induced elevation of plasma in free amino acid pool during environmental stress in the gill of
cortisol in rainbow trout (Oncorhynchus mykiss). J Exp Biol the oyster Crassostrea Virginica. Comp Biochem Physiol
205:3679–3687 71A:591–598
Li P, Gatlin DM (2006) Nucleotide nutrition in fish: current Powell MD, Ransome J, Barney M, Duijf RMM, Flik G (2007) Effect
knowledge and future applications. Aquaculture 251:141–152 of dietary inclusion of n-acetyl cysteine on mucus viscosity and
Li P, Gatlin DM (2007) Evaluation of dietary supplementation of susceptibility of rainbow trout, Oncorhynchus mykiss, and
b-hydroxy-b-methylbutyrate for hybrid striped bass Morone Atlantic salmon, Salmo salar, to amoebic gill disease. J World
chrysops 9 Morone saxatilis. J Appl Aquac 19:77–88 Aquac Soc 38:435–442
Li P, Yin Y, Li D, Kim WK, Wu G (2007) Amino acids and immune Riley WW, Higgs DA, Dosanjh BS, Eales JG (1996) Influence of
function. Br J Nutr 98:237–252 dietary arginine and glycine content on thyroid function and
Lin Y, Zhou X (2006) Dietary glutamine supplementation improves growth of juvenile rainbow trout, Oncorhynchus mykiss (Wal-
structure and function of intestine of juvenile Jian carp (Cyprinus baum). Aquac Nutr 2:235–242
carpio var. Jian). Aquaculture 256:389–394 Rubio VC, Sánchez-Vázquez FJ, Madrid JA (2006) Oral serotonin
Mai K, Zhang L, Ai Q, Duan Q, Zhang C, Li H, Wan J, Liufu Z administration affects the quantity and the quality of macronu-
(2006a) Dietary lysine requirement of juvenile seabass (Lateo- trient selection in European sea bass (Dicentrarchus labrax L.).
labrax japonicas). Aquaculture 258:535–542 Physiol Behav 87:7–15

123
New developments in fish amino acid nutrition

Saavedra M, Conceição LEC P, Pousão-Ferreira P, Dinis MT (2008) Toyohara H, Ito K, Touhata K, Kinoshita M, Kubota S, Sato K,
Metabolism of tryptophan, methionine and arginine in Diplodus Ohtsuki K, Sakaguchi M (1997) Effect of maturation on the free
sargus larvae fed rotifers: effect of amino acid supplementation. or bound forms of hydroxyproline of ayu muscle. Fish Sci
Amino Acids 35:59–64 63:843–844
Salze G, Craig SR, Schwarz M, McLean E (2008) Novel live feed Trushenski JT, Kasper CS, Kohler CC (2006) Challenges and
enrichments beneficially impact digestive ongogeny in larval opportunities in finfish nutrition. North Am J Aquac 68:122–140
cobia Rachycentron candadum. Abstract of Aquaculture Amer- Vijayan MM, Mommsen TP, Glémet HC, Moon TW (1996)
ica 08, p 337 Metabolic effects of cortisol treatment in marine teleost, the
Shamushaki VAJ, Kasumyan AO, Abedian A, Abtahi B (2007) sea raven. J Exp Biol 199:1509–1514
Behavioural responses of the Persian sturgeon (Acipenser Wilson RP (2002) Protein and amino acids. In: Halver JE, Hardy RW
persicus) juveniles to free amino acid solutions. Mar Fresh (eds) Fish Nutrition, 3rd version. Elsevier Science, San Diego,
Behav Physiol 40:219–224 USA, pp 144–179
Silvia G, Antonio UA, Francisco U, Georgina H (2002) Ammonia Wu G (1998) Intestinal mucosal amino acid catabolism. J Nutr
efflux rates and free amino acid levels in Litopenaeus vannamei 128:1249–1252
postlarvae during sudden salinity changes. Aquaculture Wu G, Bazer FW, Davis TA, Jaeger LA, Johnson GA, Kim SW,
233:573–581 Knabe DA, Meininger CJ, Spencer TE, Yin YL (2007) Important
Siwicki AK, Morand M, Fuller JC, Nissen S, Goryczko K, roles of the arginine family amino acids in swine nutrition and
Ostaszewski P, Kazun K, Glompski E (2003) Influence of production. Livest Sci 112:8–22
feeding the leucine etabolite b-hydroxy b-methyl butyrate Wu G, Bazer FW, Datta S, Johnson GA, Li P, Satterfield MC, Spencer
(HMB) on the non-specific cellular and humoral defence TE (2008) Proline metabolism in the conceptus: Implications for
mechanisms of rainbow trout (Oncorhynchus mykiss). J Appl fetal growth and development. Amino Acids. doi:
Ichthyol 19:44–48 10.1007/s00726-008-0052-7
Snyder GS, Gaylord TG, Barrows FT, Hardy RW (2008) Carnosine Wu G, Fang YZ, Yang S, Lupton JR, Turner ND (2004) Glutathione
supplementation on an all-plant protein diet for rainbow trout metabolism and its implications for health. J Nutr 134:489–492
Oncorhynchus mykiss. Abstract of Aquaculture America 08, Wu G, Morris SM Jr (1998) Arginine metabolism: nitric oxide and
p 369 beyond. Biochem J 336:1–17
Sunde J, Taranger GL, Rungruangsak-Torrissen K (2001) Digestive Wu G, Thompson JR (1989) Methionine transamination and gluta-
protease activities and free amino acids in white muscle as mine transaminases in skeletal muscle. Biochem J 262:690–691
indicators for feed conversion efficiency and growth rate in Yao K, Yin YL, Chu W, Liu Z, Deng D, Li T, Huang R, Zhang J, Tan
Atlantic salmon (Salmo salar L.). Fish Physiol Biochem 25:335– B, Wang W, Wu G (2008) Dietary arginine supplementation
345 increases mTOR signaling activity in skeletal muscle of neonatal
Szebedinszky C, Gilmour KM (2002) The buffering power of plasma pigs. J Nutr 138:867–872
in brown bullhead (Ameiurus nebulosus). Comp Biochem Yoo JH, Takeuchi T, Tagawa M, Seikai T (2000) Effect of thyroid
Physiol B 131:171–183 hormones on the stage-specific pigmentation of the Japanese
Takeuchi T (2007) Amino acids, Peptides. In: Nakagawa H, Sato M, flounder Paralichthys olivaceus. Zool Sci 17:1101–1106
Gatlin DM (eds) Dietary supplements for the health and quality Zhang Y, Dabrowski K, Hliwa P, Gomulka P (2006) Indispensable
of cultured fish. CAB International, Oxon, UK, pp 47–63 amino acid concentrations decrease in tissues of stomachless
Tapia-Salazar M, Cruz-Suarez LE, Ricque-Marie D, Pike IH, Smith fish, common carp in response to free amino acid- or peptide-
TK, Harris A, Nygard E, Opstvedt J (2004) Effect of fishmeal based diets. Amino Acids 31:165–172
made of stale or fresh herring and added crystalling biogenic Zhou X (2005) Use of synthetic lysine in fish feeds: a review on
amines on growth and survival of blue shrimp Litopenaeus research and application. Feed Ind 27:1–7
stylirostris fed practical diets. Aquaculture 242:437–453

123