World's Poultry Science Journal

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Re-evaluation of the classical dietary

arginine:lysine interaction for modern poultry
diets: a review

D. Balnave & J. Barke

To cite this article: D. Balnave & J. Barke (2002) Re-evaluation of the classical dietary
arginine:lysine interaction for modern poultry diets: a review, World's Poultry Science Journal,
58:3, 275-289, DOI: 10.1079/WPS20020021

To link to this article: https://doi.org/10.1079/WPS20020021

Published online: 23 Sep 2019.

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Reviews
Re-evaluation of the classical dietary
arginine:lysine interaction for modern
poultry diets: a review
D. BALNAVE’ and J. BRAKEZ*

‘Faculty of Veterinary Science, University of Sydney, Camden, New South Wales

2570, Australia and zDepartment of Poultry Science, College of Agriculture and
Life Sciences, North Carolina State University, Raleigh, North Carolina 27695-
7608, USA

The nutritional antagonism of arginine (Arg) and lysine (Lys) was first identified and
investigated in the 1950’s and 1960’s. The results of this early research suggest the
optimum Arg:Lys ratio to fall between 0.8 and 1.7, depending upon dietary levels of
electrolytes such as sodium, potassium, and chloride. Calculations from the more
recent optimum amino acid balances included in widely referenced authoritative
sources suggest the optimum Arg:Lys ratio to be in the range from 0.90 to 1.18.
Changes from the “optimum” value of the ratio can have adverse effects on the
performance of growing poultry. The effect is more evident with an excess of lysine
(low Arg:Lys ratio) than with an excess of arginine (high Arg:Lys ratio). Studies with
heat-stressed broilers have shown that the optimum Arg:Lys ratio varies with
ambient temperature. The Arg:Lys ratio for optimum broiler body weight gain and
feed efficiency increases at high temperatures, probably because of a reduced uptake
of arginine from the digestive tract. The improved response of broilers to increasing
dietary Arg:Lys ratio is most clearly seen during heat stress with diets containing
minimum concentrations of NaCI. This response diminishes with high dietary NaCl
and with NaHCO, supplementation, when the optimum dietary Arg:Lys ratio
declines from the high ratio (-1.30) observed with low dietary NaCl. Furthermore,
the nature of the methionine activity source influences the optimum dietary Arg:Lys
ratio for heat-stressed broilers. The performance of broilers fed 2-hydroxy-4-
(methylthio) butanoic acid (HMB) is optimised at high Arg:Lys ratios (1.35) whereas
broilers fed equimolar supplements of DL-methionine (DLM) tend to optimise
performance at lower Arg:Lys ratios (1.05). The selection of the correct methionine
activity source as a dietary supplement is likely to become more important if the
current trend to exclude animal protein feed ingredients with low Arg:Lys ratios
from poultry diets continues.

Keywords: arginine; lysine; Arg:Lys ratio; methionine; dietary electrolytes

“Corresponding author: e-mail: jbrake@ncsu.edu

0World’\ Poultry Science Aswcldtion 2002

World’s Poultry Science Journal, Vol. 58, September 2002 275
Arginine:Lysine ratio in poultry: D. Bulnave and J. Brake

Introduction: identification of the arginine:lysine (Arg:Lys) antagonism

This classic dietary antagonism was first recognised through experiments conducted with
growing chickens fed diets containing excessive lysine. These classical experiments
involved either feeding diets containing ingredients that contained high concentrations of
this essential amino acid, such as casein (Arg:Lys = 0.45), or by supplementing diets with
lysine.HC1. Many of these studies included ingredients not normally found in commercial
diets and amino acid supplements that were multiples of normal dietary inclusion levels.
However, an understanding of the role of the Arg:Lys ratio for modern poultry and poultry
diets requires an understanding of this classical literature.
Anderson and Combs (1952) reported that a depression in the body weight gain (BWG)
of chickens that resulted from feeding dietary lysine supplements of up to 40 g/kg, in
adequate or protein-limiting diets, was partially offset by the addition of gelatine (Arg:Lys
= 2.0). Anderson and Dobson (1959) later concluded that the amino acid composition of
casein was responsible for the increased arginine requirement observed when chickens
were fed diets in which casein was the primary protein source. Anderson and Dobson
(1959) obtained evidence that lysine was the amino acid most likely to influence the
arginine requirement of chickens but they concluded that other essential amino acids were
also involved. Snetsinger and Scott (1961) reported that glycine and/or arginine
supplementation of glucose-soybean meal, glucose-sesame meal and maize-soybean meal
diets containing 40 g lysine.HCl/kg helped overcome the growth depression associated
with the excess lysine. In these studies the excess lysine produced abnormal feathering
similar to that observed with chicks fed arginine-deficient diets.
Jones (1961) observed that lysine supplements of 15-20 g k g to a casein-gelatine based
diet containing 17 g lysinekg depressed BWG and caused a deterioration in feed
conversion ratio (FCR) in 1-3 week old broiler chickens. Jones (1961) reported that the
effect of the hydrochloride salt of lysine was more severe than that of the free base.
Toxicity symptoms, such as hyper-irritability, leg tremors and leg weakness, were also
evident in the lysine-supplemented birds. Excess lysine increased the lysine concentration
in plasma, liver, and leg and breast muscle. Corresponding increases in the sodium (Na),
and decreases in the potassium (K), concentrations of leg and breast muscle were observed
but no changes were noticed in the plasma concentrations of these ions. Subsequent
studies reported by Jones (1964) showed that the toxicity resulting from high
concentrations of dietary lysine was due to an Arg:Lys antagonism, a conclusion in
keeping with the observation of Anderson and Combs ( 1 952) concerning the toxicity
alleviating effect of additional gelatine, a richer source of arginine than lysine (Arg:Lys =
2.0).
Jones (1 964) used a casein-gelatine based control diet containing 16.5 g lysinekg and
13.0 g argininekg to which supplements of up to 40 g lysinekg depressed BWG while a
dietary supplement of 20 g lysine/kg increased plasma and muscle lysine concentrations
and decreased plasma and muscle arginine concentrations. There were 2-fo1d and 3-fold
increases in plasma and muscle lysine, respectively, and corresponding reductions in
plasma and muscle arginine to one-third of values obtained from birds fed the control diet.
Addition of arginine to the 20 g/kg lysine-supplemented diet at concentrations up to 15
g/kg prevented the occurrence of the symptoms of an Arg:Lys antagonism with the best
response being observed with a supplement of 5 g/kg. The addition of I5 g arginine/kg to
the 20 g lysinekg-supplemented diet increased rather than reduced the plasma and muscle
free lysine concentrations but did increase the free arginine concentrations of these tissues.
There were also 2.5-fold increases in plasma and muscle free ornithine concentrations in
chickens receiving the I5 g/kg arginine supplement.
O’Dell and Savage ( 1 966) reported that a 20 g lysine.HCl/kg supplement depressed the

276 World’s Poultry Science Journal, Vol. 58, September 2002

 Arginine:Lysine ratio in poultry: D. Balnave and J. Brake

BWG of broiler chicks fed a glucose-casein diet containing 25 g lysinekg and 18 g

argininekg while 40 g 1ysine.HClkg added to a glucose-soybean meal diet containing 17
g lysinekg and 18 g argininekg also depressed BWG. Supplementary glycine was
without effect with the former diet but arginine supplements largely overcame the growth
depression of the latter diet. By feeding a glucose-sesame meal diet containing 8 g
lysinekg and 24 g argininekg these workers found that excess arginine had a much
smaller effect on BWG than did excess lysine. These workers concluded that a metabolic
antagonism existed between arginine and lysine.
Although Boorman and Fisher (1966) argued against the existence of a specific Arg:Lys
interaction, Dean and Scott (1 968) obtained a consistent dose-related improvement in
FCR and a reversal of growth depression caused by the supplementation of a crystalline
amino acid diet with 12.5 g 1ysine.HClkg and dietary supplementation of 2.4 g to 12.1 g
arginine.HClkg. Furthermore, work reported by D’Mello and Lewis (1 970) clearly
showed the existence of a specific Arg:Lys relationship. These workers found that the
growth depression caused by the addition of lysine to a series of diets limiting in
methionine, tryptophan, histidine or threonine was reversed by supplementation with
arginine and not by the originally limiting amino acid. These workers concluded that “the
interaction between lysine and arginine cannot be shown not to be specific.” They also
observed that excess dietary lysine increased plasma lysine concentrations and reduced
plasma arginine concentrations. Addition of arginine to the diet containing excess lysine
generally, but not always, further elevated plasma lysine as well as increasing plasma
arginine concentrations. Jones et al. (1967), using crystalline amino acid diets, also noted
inconsistent responses in plasma lysine, but not plasma arginine, concentrations when
arginine was added to diets containing different concentrations of lysine.
Jones et al. (1967) observed that excess dietary lysine elevated kidney arginase activity
in chicks. Austic and Nesheim (1970) reported that a number of other essential amino
acids, including histidine, tyrosine and isoleucine also increased kidney arginase activity.
Smith (1968) had previously shown that excesses of lysine, tyrosine, histidine, glycine,
valine, threonine, cystine and methionine added to a diet limiting in arginine and
methionine induced depressions in FI and BWG that were overcome to a limited extent by
additional arginine. Austic and Nesheim (1970) concluded that alterations in the rate of
arginine degradation resulting from dietary amino acid excesses affected arginine
requirements only in diets limiting or barely adequate in arginine. They also showed that
threonine and glycine reduced the activity of arginase in the kidney of chickens fed
glucose-casein based diets first limiting in arginine. In fact, threonine reduced the activity
of this enzyme in the presence or absence of supplementary arginine. The effect of other
amino acids on arginine degradation and, therefore, on the arginine concentration
necessary to achieve the ideal dietary Arg:Lys ratio, could be extremely important in
defining arginine requirements with different feed formulations that produce different
amino acid balances.
As a result of the studies reported above, by 1970 the specific antagonism between
arginine and lysine was generally recognised as an important amino acid interaction with
ramifications for dietary formulation.

Possible mode of action of the Arg:Lys antagonism

Early suggestions concerning the nature of this amino acid antagonism included a reduced
availability of arginine from specific feed ingredients in the presence of excess lysine
(Jones, 1964) and the utilisation of arginine to aid the excretion of excess nitrogen
(Snetsinger and Scott, 1961), especially in the presence of excessive dietary essential

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Arginine:Lysine ratio in poultry: D. Balnave arid J. Bruke

amino acid concentrations. The benefit arising from a combination of glycine and arginine
in some early studies was thought to be due to the importance of glycine in aiding the
excretion of excess nitrogen via the uric acid cycle (Snetsinger and Scott, 1961). However,
Austic and Nesheim (1 970) subsequently showed that glycine depressed kidney arginase
activity in chickens. A possible increase in the rate of conversion of arginine to creatine
stimulated by excess dietary lysine was refuted by a reduced liver transamidinase activity
(Jones et al., 1967).
A suggested mechanism for the antagonism was based upon the fact that arginine and
lysine are both basic amino acids that may compete for absorption from the digestive tract.
This explanation was negated by the studies of Jones et al. (1967), who reported that
lysine did not interfere with the digestion or absorption of arginine. Jones et ul. (1967) also
showed that kidney arginase was elevated by excess dietary lysine but that the increase in
the activity of this enzyme occurred much later than the decrease in plasma arginine
which, in turn, was not associated with an increased concentration of omithine in plasma
and other tissues. It would appear that the initial problem relates to competition between
lysine and arginine for renal tubular resorption in that intravenous infusion of lysine into
cockerels at rates between 0.5 and 4.0 pmol/min/kg was shown to increase plasma lysine
concentrations and lysine excretion rates while inhibiting renal reabsorption of arginine
(Boorman et al., 1968). The reported interaction with potassium and sodium salts of
metabolisable anions (see below) may be associated with a reduced uptake of lysine by the
kidney tubules resulting from the metal cation combining with neutral amino acids to
occupy lysine binding sites (Thomas et al., 1971). The competition for renal absorption is
reflected in plasma and tissue amino acid concentrations.
Austic and Nesheim (1970) found that increased urea excretion following excess dietary
lysine addition to an arginine-limiting chick diet paralleled changes in renal arginase
activity and these correlated closely with reductions in FI and BWG. They also showed
that kidney arginase in chicks was increased by supplementing a glucose-casein diet with
either 4 g lysine.HCl/kg or 10 g arginine.HCl/kg although both the BWG and the FCR
resulting from these treatments varied in opposing directions relative to the basal diet.
They reported that chickens with elevated kidney arginase activity had characteristically
elevated plasma lysine concentrations.

Electrolyte influences on dietary Arg:Lys antagonism

Jones (1961) originally reported that excess lysine added to a casein-gelatine based diet
induced increases in the sodium and decreases in the potassium concentrations in leg and
breast muscle but not in plasma. In subsequent studies Jones (1 964) again noted similar
changes in cellular sodium and potassium concentrations as a result of excess lysine
supplementation of casein-gelatine and soy-protein diets. These observations suggested
that an interaction existed between electrolytes and dietary Arg:Lys ratio.
O’Dell and Savage (1966) examined the effect of dietary supplements of potassium
acetate (KAc) and potassium chloride (KCI) on the Arg:Lys antagonism induced in
glucose-casein, glucose-soybean meal protein and glucose-sesame meal based diets.
Some interesting effects were evident in these data. In the case of glucose-casein based
diets, supplementation with 27 g KAc/kg increased BWG in the presence or absence of
additional lysine (Arg:Lys ratios of 0.29 and 0.48, respectively). With the glucose-soybean
meal based diet, 27 g KAc/kg gave improved BWG in the presence of additional lysine
(Arg:Lys of 0.38) but not with the basal diet (Arg:Lys of 1.06). In the case of the high-
arginine, glucose-sesame meal based diet, BWG to 4 weeks of age was improved with
both 27 g KAckg and 20 g KClkg to an optimum dietary Arg:Lys ratio of 1.7. However,

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at higher Arg:Lys ratios, BWG was superior with the 20 g KCl/kg addition whereas 27 g
KAckg supplementation gave better BWG at lower Arg:Lys ratios. Increasing the dietary
Arg:Lys ratio from 2.0 to 3.0 through supplementation with arginine.HC1 reduced BWG
in the presence of 27 g KAcIkg but not with 20 g KClkg. Within the dietary Arg:Lys ratio
range from 1.09 to 1.64 neither potassium salt produced a significant difference in BWG.
These data indicate that KAc was most effective at low (<1.0) dietary Arg:Lys ratios
suggesting that KAc ameliorates the effects of excess lysine. In contrast, KCI was most
effective at high (>I .7) dietary Arg:Lys ratios suggesting that chloride ameliorates the
effects of excess arginine. It is possible that these effects were induced by changes in the
digestibility or catabolism of arginine and lysine, or other amino acids, but this evidence
is lacking at present.
Stutz et al. (197 1) used a casein based diet with an Arg:Lys ratio of 0.48, similar to that
used by O’Dell and Savage (1966). They reported improved BWG with dietary additions
of arginine.HC1, KAc, NaAc and potassium carbonate (K,CO,) but not with KCl. Again,
the KAc and NaAc appeared to ameliorate the effect of excess dietary lysine. By studying
the effects of additional arginine.HC1, KAc and K,CO, on the alleviation of ataxic gait,
abnormal feathering and depressed BWG, all indicators of arginine deficiency, these
workers suggested that with a diet containing excess lysine an excess of dietary cations,
from salts of metabolisable anions, had a beneficial effect on broiler performance. They
hypothesised that excess cations decrease the rate of catabolism of arginine by preventing
the induction of kidney arginase. This suggestion received qualified support in a
subsequent study (Stutz et al., 1972) although the response of chickens fed KAc was
inconsistent.
Stutz et a/. (1971) found the concentrations of free lysine in the plasma and breast
muscle of broilers fed a high-lysine casein based diet were reduced by feeding additional
KAc alone or with arginine.HC1. The free lysine concentration in plasma, but not breast
muscle, was reduced by feeding additional arginine.HC1. The feeding of additional
arginine.HC1 and KAc, separately or in combination, increased the concentrations of free
arginine in plasma and breast muscle. These responses in plasma amino acid
concentrations were confirmed in a subsequent study where supplements of arginine.HC1,
KAc, K,CO, or NaAc to a diet with an Arg:Lys ratio of 0.72 all reduced the lysine
concentration, while all but NaAc increased the arginine concentration, in plasma. The
concentration of free ornithine in the plasma was elevated with the arginine supplement
only and reductions were observed with the K,CO, and NaAc additions.
However, Scott and Austic (1978) concluded that high concentrations of cations,
including potassium, in diets containing excess lysine increased the catabolism of lysine
so that a more favourable dietary Arg:Lys ratio was achieved. Interestingly, a similar
response was found with dietary addition of arginine. In studies using a similar basal diet,
Stutz et al. ( 1972) found arginine.HC1 and KAc supplements reduced plasma and kidney
lysine, and increased plasma and kidney arginine, concentrations but only treatments
involving arginine.HC1 increased ornithine concentrations. Stutz et ul. ( 1972) concluded
that KAc supplementation of a glucose-casein based diet (low Arg:Lys ratio) decreased the
activity of kidney arginase and the resulting increased availability of arginine allowed
increased use of all amino acids for protein synthesis.
In contrast to the results of Stutz et al. (1971, 1972), Scott and Austic (1978) failed to
observe any effect of KAc supplementation on kidney arginase activity although
arginine.HC1 did significantly increase the activity of this enzyme. Also, in agreement
with previous work (Austic and Neshiem, 1970; Stutz et al., 1971, 1972), they reported
that dietary supplementation with arginine.HC1 reduced the plasma concentration of
lysine and increased the plasma concentration of arginine. Similar responses were
observed with a dietary supplement of KAc (Scott and Austic, 1978). These plasma lysine

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Arginine:Lysine ratio in poultry: D. Balnave and J. Brake

responses to additional dietary arginine contrasted with those reported by Jones (1964)
and D’Mello and Lewis (1 970) where dietary arginine supplementation increased plasma
lysine concentrations. These and other studies (Jones et al., 1967) highlight the
inconsistent responses in plasma lysine concentrations to dietary arginine
supplementation.
The influence of dietary chloride concentration on the responses of growing egg-type
chickens to changes in dietary Arg:Lys ratio was examined by Calvert and Austic (198 1).
Increasing concentrations of dietary chloride as calcium chloride (CaC1,) from 4.4 to 18.4
g k g had no significant influence on plasma lysine or arginine in glucose-maize gluten
meal based diets with Arg:Lys ratios of I .23 and 1.92 although BWG and feed intake (FI)
were reduced and FCR was increased at higher concentrations of CaC1,. With diets
formulated from similar ingredients, and with Arg:Lys ratios of 0.40, 0.83 and 1.43 at a
similar dietary arginine concentration of 10 g/kg, increasing the dietary chloride
concentration (as CaCl, and HCI) from 5 to 15 g k g induced reductions in BWG, and at an
Arg:Lys ratio of 0.40 an increase in FCR, at higher chloride concentrations. Overall mean
BWG was maximised with the lysine-adequate diet with an Arg:Lys ratio of 0.83 and
miniinised with the excess lysine diet with an Arg:Lys ratio of 0.40. Increasing the dietary
arginine concentration in the diet containing excess lysine to adjust the Arg:Lys ratio from
0.40 to 0.92 improved BWG to that observed with the lysine adequate diet containing an
Arg:Lys ratio of 0.83 although BWG still declined with increasing dietary chloride
concentration. These workers concluded that excess dietary chloride increased the degree
of antagonism between arginine and lysine. Subsequently, Austic et al. (1986) suggested
that dietary electrolyte balance was important with values below 100 mEqkg of diet
markedly exacerbating an Arg: Lys antagonism induced by excess dietary lysine.
Supplementing a diet with cationic salts of metabolisable anions elevates the dietary
electrolyte balance, a response not obtained with neutral salts.

General conclusions from classical studies

Although derived from studies with non-practical diets with often excessive
concentrations of amino acids, the classical data do indicate that the optimum Arg:Lys
ratio falls somewhere between 0.8 and 1.7 depending on the dietary Na, K and C1
concentrations, i.e. dietary electrolyte balance.
Table 1 Suggested argininedysine ratios for broilers derived from various authorities.

Age Arginine Lysine Arginine:lysine Authority

(weeks) % 8 ratio

0-3 1.25 1.10 1.14 National Research

3-6 1.10 1.00 1.10 Council, USA (1 994)
6-8 1.00 0.85 1.18

0-4 1.02 1.13 0.90 Standing Committee

4-8 0.94 0.90 I .04 on Agriculture,
Australia (1987)

0-4 I .03 1.10 0.94 Agricultural Research

4-8 0.76 0.80 0.95 Council, UK (1 975)

0-3 1.18 1.12 1.05 Baker (1997)

3-6 0.96 0.89 1 .08
6-8 0.82 0.76 1 .ox

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 Arginine:Lysine ratio in poultry: D. Balnave and J. Brake

These conclusions need to be examined in the light of modern genetics and practical
dietary formulations. Calculation of the Arg:Lys ratio derived from the estimated amino
acid requirements produced by various authorities are shown in Table 1. Those produced
by the Agricultural Research Council (1973, Australian Standing Committee on
Agriculture (1987), and National Research Council (1994) are derived from a review of
the literature and suggested ratios range between 0.90 and I . 18. Those derived from the
experimental data of Baker and associates from the University of Illinois range between
1 .05 and 1.08. These latter experiments used a standard crystalline diet containing 8.8 g
NaClkg (Baker et al., 1979). This diet contains a high concentration of NaCl compared to
conventionally formulated diets.

Identification of the need for increased dietary Arg:Lys ratios during heat
stress
The studies reported in this section vary to some extent from those reported above in that
they mainly involve the feeding of diets composed of conventional feed ingredients and
containing conventional concentrations of lysine, to which supplements of arginine free
base were used to vary the dietary Arg:Lys ratio.
Brake et al. (1994ab; 1998) drew attention to the need for dietary Arg:Lys ratios to be
increased above generally recognised values in order to optimise the performance of heat-
stressed broilers exposed to temperatures of approximately 3 1"C between 2 1 and 49 days
of age. Brake et al. (1994ab; 1998) first reported that dietary Arg:Lys ratios of
approximately 1.35 appeared optimal compared to the reference ratios shown in Table I .
The most consistent response obtained in the initial studies was an improvement in FCR
without any loss in BWG. This response was obtained either when arginine free base was
used as a dietary supplement in place of an inert filler or when practical diets with
differing feed ingredients were used to vary the Arg:Lys ratio. The need for additional
arginine during heat stress appeared to be related to a reduced uptake of' arginine from the
digestive tract of broilers since in vitro epithelial uptake studies showed that in the
presence of an equimolar concentration of lysine at 31°C the uptake of arginine was
reduced significantly compared with the uptake at 22°C.
Studies by Gorman et a1. (1997) showed that the dietary Arg:Lys ratio could influence
the breast meat yield of broilers. Increasing the dietary Arg:Lys ratio increased breast meat
yield at moderate to cool temperatures but reduced breast meat yield at high temperatures.
However, the dietary Arg:Lys ratio required to optimise breast meat yield at the cooler
temperatures was found to be higher than the Arg:Lys ratio calculated from NRC (1994)
recommendations.
Inconsistent results have been reported by others. Mahmoud and Teeter (1996) fed diets
containing Arg:Lys ratios of 1.1 and 1.4 to broilers at thermoneutral (24°C) and cycling
heat stress (24-35°C) temperatures and failed to obtain any significant effect of Arg:Lys
ratio on production at 30 and 42 days of age or any Arg:Lys x temperature interactions on
production or carcass composition. However, increasing the Arg:Lys ratio significantly
increased dressing percentage and breast meat yield. Mendes et al. ( I 997) supplemented a
basal diet containing 10 g lysinekg, 11 g argininekg and 3.3 g NaClkg with both lysine
and arginine to obtain diets with 10, 11 and 12 g lysine/kg and Arg:Lys ratios of 1.1, 1.2,
1.3 and 1.4 and fed these diets to 21-42 day old broilers maintained at either a constant
15.5"C, a constant 21.1"C or a cycling 25.5-33.3"C. They found a linear (P=O.101)
increase in breast meat yield coupled with a significant (P=0.006) linear reduction due to
the Arg:Lys ratio in abdominal fat content across all thermal environments. They also
reported that increasing the Arg:Lys ratio significantly improved FCR (up to a ratio of 1.3

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over all thermal environments) and no significant Arg:Lys by temperature interaction was
observed. At the hot temperatures the FCR at Arg:Lys ratios of 1.3 and 1.4 (1.91 6 and
1.912) were considerably improved compared with Arg:Lys ratios of 1.1 and 1.2 ( I .9S5
and 1.961). Similar, and significant, changes in FCR with increasing dietary Arg:Lys ratio
were obtained by Brake et al. (1998) using 28-49 day-old broilers maintained at a constant
3 1°C and fed diets containing 10.2 g lysine/kg with a dietary NaCl supplement of 1.2 g k g .
The FCR values were 2.383 and 2.294 at Arg:Lys ratios of 1.OS and 1.20, and 2.165 and
2.022 at Arg:Lys ratios of 1.34 and 1.49. Taken together, the data shown above suggest that
in the case of heat stressed broilers, performance will be improved by the use of dietary
Arg:Lys ratios in excess of 1.2, depending on the dietary concentration of NaCl.

Electrolyte effects on optimum dietary Arg:Lys ratio during heat stress

Brake et ul. ( I 998) reported that increasing the Arg:Lys ratio of diets fed to Australian
broilers at 3 1 “C by using additional arginine free base gave significant improvements in
BWG and FCR when the diets contained 1.2 g NaClkg but not when the diets contained
2.4 g NaClkg. In fact, the optimum dietary Arg:Lys ratio for BWG and FCR decreased
from 1.34 with the lower NaCl supplement to 1 .OS with the higher NaCl supplement. The
additional NaCl gave significantly better FCR at an Arg:Lys ratio of 1.05 and a
significantly poorer FCR at an Arg:Lys ratio of 1.49. A possible explanation, based upon
data discussed above, is that the increase in the sodium cation concentration reduced
lysine resorption by the kidney tubules thereby producing an “effective” Arg:Lys ratio
higher than suggested by the calculated 1 .OS ratio (Thomas et al., 1971). This may have
been beneficial with the calculated 1.05 ratio diet but may have caused an excessively high
ratio with the calculated 1.49 ratio diet. Alternatively, the additional cations could have
contributed to lysine catabolism (Scott and Austic, 1978) with the same result. The fact
that the results of O’Dell and Savage (1 966) suggest that chloride may ameliorate the
effects of excess arginine, thereby reducing the “effective” Arg:Lys ratio, is a further
nutritional complication. Under USA conditions, Brake et al. (1998) included a dietary
NaCl supplement of 3.3 g/kg, similar to that used by Mendes et ul. (l997), in diets for 42-
5.5 day-old heat stressed and thermoneutral broilers and found a significant thermal
environment x Arg:Lys ratio interaction for FCR due to a linear improvement in FCR in
heat stress, but not thermoneutral, conditions.
Balnave and Brake (1999) examined the use of a dietary supplement of 16 g sodium
bicarbonate (NaHCOJkg instead of the supplement of 1.2 g NaClkg using the same
basal diet described by Brake ef al. ( I 998). Diets containing Arg:Lys ratios ranging from
1.05 to I .3S were fed to Australian broilers at 2 1 “C and 3 1 “C and the Arg:Lys ratios were
varied using arginine free base. The responses were not as clear cut as with the NaCl study
with trends rather than significant differences being observed. Nevertheless, at 3 1“C there
was a consistent but non-significant improvement in F1 and BWG with increasing dietary
Arg:Lys ratio in the absence of NaHCO,. Supplementation with NaHCO, improved FI and
BWG at Arg:Lys ratios of 1.15 and I .2S. This indicated that at 3 1 “C the optimum dietary
Arg:Lys ratio in the presence of this concentration of NaHCO, was in the range I . 1 S- 1.25.
Balnave and Brake ( 1999) also found that increasing the dietary ArgLys ratio through
arginine supplementation resulted in increases in plasma arginine and ornithine without
any consistent effect on plasma lysine. However, a slower rate of increase in plasma
ornithine with increasing dietary Arg:Lys ratio was observed at 31°C than at 21”C, and
lower plasma concentrations of ornithine were observed at the higher temperature at each
dietary Arg:Lys ratio, implying less degradation and, therefore, greater utilisation of
arginine at 31°C. The only treatment to show an increase in plasma Arg:Lys ratio in

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response to the maximum dietary Arg:Lys ratio of 1.35 was the NaHCO, treatment at
31°C. This indicated the NaHCO, to be important in potentiating the elevated Arg:Lys
ratio in plasma. These authors suggested that, at heat stress temperatures, NaHCO, may
act by supplying bicarbonate ions to buffer the guanidinium group of arginine, a more
strongly charged cation than lysine, in the blood. Maybe cationic salts of metabolisable
anions such as NaHCO, act by buffering the arginine once it is absorbed into the blood.
Separating the mechanism of action of NaHCO, from the digestive tract is in keeping with
the fact that earlier studies had shown that NaHCO, had no effect on the ileal digestibility
of lysine or arginine (Balnave and Oliva, 1991).
In a subsequent study Balnave and Brake (2001) fed diets with or without a 16 g
NaHCO,/kg supplement to broilers maintained in either a constant 20"C, a constant 3 1°C
or a diurnal cycling 2535°C environment. Practical dietary formulations using different
feed ingredients were used with one diet containing only animal-sourced, protein-rich
ingredients (Arg:Lys= 1.01) and the other diet containing predominantly plant-sourced,
protein-rich ingredients (Arg:Lys=l.31). Supplementation with NaHCO, had no
significant overall effect on FI, BWG or FCR but significant NaHCO, by temperature
and NaHCO, by diet interactions were observed for BWG. Sodium bicarbonate
improved BWG at constant 31°C but not at constant 20°C or in the diurnal temperature
environment. Sodium bicarbonate significantly improved BWG with the diet containing
an Arg:Lys ratio of 1.31 but not with the diet containing an Arg:Lys ratio of 1.01. These
results agreed with those reported in earlier work (Balnave and Brake, 1999) and taken
together suggest that for optimum performance the highest Arg:Lys ratio for heat-stressed
broilers receiving NaHCO, at 3 1 "C is approximately 1.30. Significant temperature by
Arg:Lys ratio interactions showed that in this study Fl and BWG were significantly
improved at the higher Arg:Lys ratio in all three temperature environments. An
improvement in BWG at thermoneutral temperatures has also been noticed on other
occasions (Brake et al., 1998).
These, and the results of earlier studies using high dietary Arg:Lys ratios reported by
Balnave and Oliva (1991), suggest that the major production responses of heat-stressed
broilers to dietary NaHCO, supplementation are manifested through improvements in FI
and BWG although an improvement in FCR is sometimes observed (Gorman and
Balnave, 1994). In the absence of NaHCO, the most consistent response observed with
heat-stressed broilers fed diets with a high Arg:Lys ratio is an improvement in FCR
although an improvement in BWG is sometimes observed (Brake et al., 1998).

Low-protein diets for heat-stressedbroilers

Recommendations for reducing the adverse effects of high ambient temperature on the FI
and BWG of broilers include reducing dietary crude protein concentrations and
supplementing the diet with individual essential amino acids (Waldroup er a/., 1976;
Waldroup, 1982; Austic, 1985). However, MacLeod (1997) found no evidence of
enhanced thermogenesis related to amino acid imbalance and the present authors have
field evidence of good production using high protein diets. Therefore, the results of studies
such as that reported by Alleman and Leclercq ( 1 997) are worth considering. In this work
the dietary protein concentration for 23-44 day old broilers at 22" and 32°C was reduced
from 200 to 160 g k g and the low-protein diet was supplemented with a range of amino
acids including lysine and arginine. No advantages were observed from feeding the amino
acid-supplemented, low-protein diet at either temperature. The adverse responses
observed at 32°C in this work were probably related to the fact that the Arg:Lys ratio
actually declined from a calculated 1.24 ratio in the 200 g k g protein diet to I .07 in the 160

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Arginine:Lysine ratio in poultry: D.Balnave and J. Brake

g k g protein diet with correspondingly significantly inferior BWG and FCR, and
numerically inferior FI, with the amino acid-supplemented, low-protein diet. In this
regard, Chen et al. (2000) reported that the removal of the lysine or lysine and methionine
supplements from a low-protein broiler finisher diet containing I66 g crude proteinkg had
no adverse effect on the performance of heat-stressed broilers even though the lysine
concentration was reduced to 7.7 g k g compared with the recommended 8.5 g k g
(National Research Council, 1994). This was presumably related to the fact that the
dietary Arg:Lys ratio was increased from 1.18 to I .30 through the removal of the lysine
supplement.

Studies with turkeys

As with broilers, research studies with turkeys have led to conflicting results. Brake et al.
(1994b) used British United Turkey (BUT) turkey toms reared in open houses in summer
in North Carolina and reported that increasing the dietary Arg:Lys ratio from 1 .05 to 1.30
between 8 and 20 weeks of age produced beneficial responses in liveability, body weight
and FCR at 20 weeks of age, in a similar fashion to that observed with heat-stressed
broilers. Higher breast meat yield was also obtained in this study. Kidd and Kerr (1998),
using BUT turkey toms housed in an enclosed power-ventilated house where the average
temperature range was 15-19"C, reported similar improvements in body weight and breast
meat yield when the dietary Arg:Lys ratio was increased from 0.98 to 1.22 between 8 and
20 weeks of age. At these temperatures no effect was noted on liveability and the
responses to increased dietary Arg:Lys ratio were not affected by increased dietary
electrolyte balance (DEB) adjusted by supplementation with NaHCO,.
Waldroup et al. ( 1 998) investigated the responses of varying Arg:Lys ratios (1 .O-1.3) at
lysine values of 100, 110 and 120% of NRC (1 994) recommendations fed to 18 weeks of
age to BUT and Nicholas 700 toms in a curtain-sided house in Arkansas: season unknown.
They reported a significant strain x Arg:Lys interaction for 18 week body weight that was
due to an increasing body weight with increasing Arg:Lys ratio in the BUT strain and the
opposite effect in the Nicholas strain. The data also showed that the BUT strain had a
slightly higher lysine requirement for maximum growth. Therefore, given the proposition
that there exists an optimum Arg:lys ratio for a given strain and environment it is
reasonable that the BUT strain would also require more arginine compared to the Nicholas
strain that performs better at a lower dietary lysine concentration. In the Nicholas strain an
equivalent Arg:Lys ratio would represent an excess of arginine and, therefore, an Arg:Lys
imbalance. It is of interest that there was an Arg:Lys ratio x strain interaction for breast
meat yield that approached significance (P < 0.10).It should be noted that the dietary NaCl
level in this experiment was higher than all the other reports cited in this section, which
should result in a decrease in the optimum Arg:Lys ratio in a manner reported for broilers
by Brake et al. (1998).
More recently, Veldkamp et al. (2000) grew male BUT birds on diets containing Arg:Lys
ratios approximating 1.O and 1.25, and DEB values approximating 164 and 254 meqkg
from 28 to 140 days of age at constant 15" and 30°C. The higher Arg:Lys ratio improved
FI to 70 days of age, and BWG and FCR to 98 days of age but not beyond. These data
suggest an acclimation effect to constant temperature regimes that were not used in the
other studies reported in this section. Nevertheless, there were some cumulative
interactions of dietary Arg:Lys ratio and DEB. A higher Arg:Lys ratio improved BWG and
FCR at the low DEB but had no effect at the higher DEB. There was no effect of treatment
on breast meat yield. It is of interest that in this study the DEB was controlled by the
addition of variable amounts of NaHCO, and ammonium chloride with no added NaCl

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 Arginine:Lysine ratio in poultry: D.Balnave and J. Brake

whereas the other studies referred to in this section used diets with a consistent supplement
of NaC1.

Effect of methionine activity sources on optimum arginine:lysine ratio

Studies to determine the relative efficacies of DL-methionine (DLM) and various
methionine hydroxy analogues, including 2-hydroxy-4-(methylthio) butanoic acid
(HMB), have produced controversial and conflicting results. Recent examination of the
diets fed in numerous studies indicated that the dietary Arg:Lys ratio may have influenced
the determined efficacy values in these early studies (see Balnave et al., 1999). Crystalline
amino acid diets, which are characterised by low Arg:Lys ratios of approximately 1 .O to
I .05 and high NaCl concentrations of 4.0 to 9.0 g k g typically show a higher efficacy for
DLM whereas differences between methionine activity sources are small with practical or
semi-purified diets containing intact proteins that typically have Arg:Lys ratios of 1.05 to
1.20 and lower NaCl concentrations. The fact that the efficacy of methionine activity
source may be linked to the dietary Arg:Lys ratio suggests that the optimum Arg:Lys ratio
during heat stress may be dependent upon the source of methionine activity. This
hypothesis was confirmed by comparing DLM and HMB at assumed equimolar
equivalence in diets varying in Arg:Lys ratio.
Balnave et al. (1 999) showed that the presence of equimolar supplements of DLM or
HMB affected the optimum dietary Arg:Lys ratio of chronically heat-stressed broilers fed
diets containing 3.0 g NaClkg. The primary response appeared to be related to a relatively
lower FI in broilers fed DLM at dietary Arg:Lys ratios 1.20 and 1.34 compared to 1.03.
Thus, in the pre3ence of HMB broilers performed similarly at Arg:Lys ratios of 1.03, 1.20
and 1.34 whereas broilers fed DLM performed best with an Arg:Lys ratio of 1.03.
Our unpublished work has included an examination of alternating periods of heat-stress
and thermoneutral temperatures. Broilers were exposed to 22°C from 28 to 33 and from 38
to 43 days of age, and to 30°C from 33 to 38 days of age. They were fed diets containing
Arg:Lys ratios of 1.04, 1.19 and 1.35 with 2.0 g NaClkg and equimolar supplements of
either DLM (2.3 g/kg) or HMB (2.6 g k g ) . The dietary Arg:Lys ratio only affected the
performance of the broilers during the 32°C exposure period. At this temperature the FI,
BWG and FCR with HMB were optimised at a dietary Arg:Lys ratio of 1.35 whereas
broilers fed DLM tended to optimise BWG and FCR at an Arg:Lys ratio of 1.04. In
agreement with the results reported by Balnave et al. (1999), FI appeared to exert a major
effect on performance. At 30°C broilers fed HMB grew significantly better than broilers
fed DLM at an Arg:Lys ratio of 1.35.
The results of these studies indicate that the optimum dietary Arg:Lys ratio for heat-
stressed broilers is dependent on the dietary methionine activity source. One possible
mechanistic basis for an interaction of dietary arginine and methionine lies in the fact that
these two amino acids, in combination with glycine, form creatine (Almquist et al., 194 I).
Arginine and glycine form glycocyamine to which is normally added a methyl group from
methionine (Keshavarz and Fuller, 1971). In this regard it is worth noting that Smith
(1968) reported that methionine was one of several amino acids that depressed FI and BW
gain when added in excess in an arginine-limiting casein-based diet. These adverse effects
could be reduced by additional arginine, implying that methionine has some effect on the
degree of antagonism between arginine and lysine, that there is some direct interaction
between arginine and methionine, or that arginine has some over-riding effect on the
metabolism of several amino acids. Amelioration of reduced growth rate due to excess
methionine by additional dietary arginine has been reported to be accompanied by
increased excretion of creatine in rats (Brown and Allison, 1948). On the other hand,

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growth depression caused by excess dietary arginine can be ameliorated by additional

dietary methionine, which is also accompanied by increased excretion of creatine (Fuller
et al., 1967; Keshavarz and Fuller, 1971). Thus, the data suggest that imbalances of
arginine and methionine can be resolved by excretion of creatine. In the case of heat stress,
there is a decrease in the intestinal uptake of arginine (Brake et al., 1994a; 1998) and
probably a decline in the requirement for methionine for growth. This would be expected
to create a situation in the circulation of relative amino acid imbalance where the supply
of arginine has declined relative to methionine, at least in the short term. Based upon the
studies outlined above, the excess methionine might be expected to be degraded more
rapidly in a diet with a narrow Arg:Lys ratio (relative deficit of arginine) while a wide
Arg:Lys ratio should support better growth. This seems to be the case with HMB but not
DLM in acute heat stress. This may be related to the fact that HMB is not absorbed as an
amino acid but is converted in the liver, possibly only as needed.

Other functions of arginine

Arginine is the precursor of the biological dilator nitrogen oxide (NO) and, as such, has
been evaluated as a preventive treatment for ascites (pulmonary hypertension syndrome).
Wideman et al. ( I 995) reported that dietary L-arginine supplementation reduced the
incidence of mortality from ascites in broilers exposed to cool temperatures of 10°-15”C,
without affecting BWG. Subsequently, Wideman et al. (1996) reported that supplementing
the diet of 38-49 day old broilers with L-arginine HCI to increase the dietary Arg:Lys ratio
from 1.18 to 1.78 allowed these birds to exhibit highly effective pulmonary vasodilation
in response to an acute increase in blood flow. This did not occur with broilers fed the
unsupplemented, low Arg:Lys ratio diet, indicating that birds fed this diet were unable to
maintain sufficiently high rates of NO production. The unsupplemented diet had lysine
and arginine concentrations of 13.3 g k g and 15.7 g k g , respectively, which were in excess
of National Research Council (1994) requirements but gave an Arg:Lys ratio of I .I 8,
similar to the ratio calculated from National Research Council (1 994) lysine and arginine
requirements.
Arginine is also known to influence immunity and disease resistance. Kwak et al. (1 999)
reported that an arginine-deficient diet had adverse effects on the lymphoid organs with
the thymus and spleen weights being more affected than bursa weight. In fact, the relative
bursa weight (as a percent of body weight) was not affected by the arginine deficiency in
this study. Also, Qureshi et al. (2000) have reported that during heat stress phagocytic
functions of monocytes and macrophages are influenced by the dietary Arg:Lys ratio. It is
well known that NO is required for optimal phagocytic function.

Conclusions
A specific relationship between dietary arginine and lysine has been shown to exist in
broilers such that changes from an “optimum” value of the ratio has an adverse effect on
BWG and FCR with a concomitant effect on plasma and muscle amino acid
concentrations. The effect is more evident with an excess of lysine (low Arg:Lys ratio)
than with an excess of arginine (high Arg:Lys ratio).
The adverse effects of an excess of lysine on BWG and FCR, and on the induction of
high plasma lysine concentrations, are ameliorated by dietary supplementation with
arginine or with certain metabolisable salts of sodium and potassium. These treatments
also improve plasma arginine concentrations. Excess lysine has been reported not to

286 World’s Poultry Science Journal, Vol. 58, September 2002

 Arginine:Lysine ratio in poultry: D. Balnave and J. Brake

interfere with the digestibility or absorption of arginine but to inhibit renal reabsorption of
arginine and to induce kidney arginase activity, the latter effect resulting in elevated
plasma lysine concentrations. While potassium appears to ameliorate the effects of excess
lysine, chloride appears to ameliorate the effects of excess arginine. Specific studies with
heat-stressed broilers, in which increases in dietary arginine were used to increase the
dietary Arg:Lys ratio, at specific lysine concentrations, have shown that the optimum
Arg:Lys ratio varies with ambient temperature. The Arg:Lys ratio for optimum BWG and
FCR increases at high temperatures, probably because of a reduced uptake of arginine
from the digestive tract.
Increases in dietary arginine increase plasma arginine and ornithine concentrations
without any consistent effect on plasma lysine. The dietary arginine concentration needed
to obtain the optimum dietary Arg:Lys ratio at a particular lysine concentration is
intluenced by dietary electrolytes, such as sodium, potassium and chloride, and by other
dietary amino acids. The improved response of broilers to increasing dietary Arg:Lys ratio
is most clearly seen during heat stress with diets containing minimum concentrations of
NaC1. The effect diminishes with high dietary NaCl and with NaHCO, supplementation,
when the optimum dietary Arg:Lys ratio declines from the higher ratio observed with low
dietary NaC1. One possibility is that the sodium cation may reduce lysine resorption by the
kidney tubules thereby inducing a higher “effective” dietary Arg:Lys ratio.
Furthermore, the nature of the methionine activity source influences the optimum
dietary Arg:Lys ratio for heat-stressed broilers. The performance of broilers fed HMB i s
optimised at high Arg:Lys ratios (1 3 5 ) whereas broilers fed equimolar supplements of
DLM tend to optimise performance at lower Arg:Lys ratios (1.05). The major factor
affecting the performance of broilers fed these methionine-activity sources is FI. The
selection of the correct methionine activity source as a dietary supplement is likely to
become more important if the current trend to exclude animal protein feed ingredients
from poultry diets continues.
The role of arginine as a donor for the production of NO also increases the importance
of this amino acid as supplemental dietary arginine has been shown to impact the
incidence of ascites as well as influence phagocyte function during heat stress. This is due
to the roles of NO in vasodilation and killing by immune cells.

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