http://www.jstage.jst.go.

jp/browse/jpsa
doi:+*.,+.+/jpsa..0.+03
Copyright ,**3, Japan Poultry Science Association.

Review

Chaiyapoom Bunchasak
Faculty of Agriculture, Kasetsart University, Bangkok, Thailand

Synthetic DL-methionine (DLM) supplements poultry diets to enhance production. The bioe$cacy of liquid methionine
is generally lower than that of powder methionine; however, if the level of total sulfur amino acids (TSAA) is set to the
commercial recommendation, the bioe$cacy of liquid methionine seems to be equal to that of powder methionine (equimolar
basis). Absorption and transportation in the segment of the jejunum di#er between liquid and powder methionine because
multiple systems are involved. Methionine supplementation in a low-protein diet alleviates the negative e#ects of heat stress.
The supplementation improves the amino acid balance and consequently promotes growth performance by enhancing feed
e$ciency, increases protein synthesis and decreases fat synthesis. Methionine supplementation also improves the immune
response through direct e#ects (protein synthesis and breakdown) and indirect e#ects (derivatives of methionine). As various
factors inuence the methionine requirement, the requirements of commercial strains are higher than those recommended by
NRC (+33.). Moreover, the methionine requirement expressed as a percentage of diet declines during the starter and grower
phases, while the requirement related to lysine is little changed (tends to increase).
Key words: growth, heat stress, immune, methionine, poultry

J. Poult. Sci., .0: +03+13, ,**3

Introduction
The diet of poultry is supplemented with synthetic
amino acids to enhance production. Wallis (+333) describes several benets of amino acid supplementation as
follows: +) reducing cost in production, ,) producing the
optimal balance of essential amino acids that presumably
enhances growth, maximizes edible meat yield, reduces
carcass fat and enables precise selection of progeny for
breeding programs, and -) balancing an animals nutrient
intake to conserve resources and minimize wastes.
Methionine is an essential amino acid for animals, particularly in poultry. In avian species, methionine is classied as a rst limiting amino acid because it is limited in
plant protein sources and because there is a strong requirement for it to support feather growth and protein synthesis. Methionine plays several roles in poultry: rst as an
essential amino acid for protein synthesis, second as a
methyl donor group for normal cellular metabolism and
for normal formation of co-enzyme S-adenosylmethionine, third as a precursor of important intermediates in
metabolic pathways such as cystine or carnitine, fourth as
an amino acid involved in polyamine synthesis, and fth as
a sulfur donor.
Deciency in methionine consumption has a signicant
Received: November +., ,**2, Accepted: January ,1, ,**3
Correspondence: Dr. C. Bunchasak, Department of Animal Science,
Faculty of Agriculture, Kasetsart University, /* Phaholyothin Rd.,
Jatujak, Bangkok +*3**, Thailand. (E-mail: agrchb@ku.ac.th)

negative impact on animals such as growth inhibition, the

induction of metabolic disorder and the reduction of
disease defensive potential. The addition of methionine to
the poultry diet has been correlated with the tendency to
have less total body fat (Rostagno et al., +33/), to improve growth performance and to reduce odor-related
compounds in excreta (Chavez et al., ,**.). On the other
hand, feeding excess dietary methionine has been reported
to impair body weight gain (Harper et al., +31*), although
Han and Baker (+33-) demonstrated that *./ excesses
of methionine are not harmful to young broiler chicks fed
corn-soybean meal diets.
Methionine is contained in both animal and plant protein sources. The need for a plant protein source has
dramatically increased recently due to the decline in sh
meal and the prohibition on the use of animal protein
sources (meat and bone meal) due to the risk of
pathogenic and toxic substance contamination (Chadd
et al., ,**,). In addition, the use of byproducts of seed oil
as a feed ingredient has increased. Therefore, more
methionine supplementation is required. Methionine
sources are used in two forms (powder and liquid). This
article discusses the e#ects of methionine supplementation
and the di#erences between the e#ects of these two
methionine sources.
Source of Methionine
Unlike many amino acids derived from fermentation
processes, DL-methionine (DLM) is produced from a

J. Poult. Sci., .0 (-)

170

complex chemical synthetic process, and the starting material for its production is acrolein (a --carbon aldehyde)
derived from propylene (a petroleum derivative)
(Aldrich, ,**1). There is no di#erence in e#ectiveness
between L-methionine and DLM (Dilger and Baker,
,**1). Industrially, the powder and liquid forms of
methionine sources are mainly used. It is known as
DL-methionine (DLM: powder form) and DL-,hydroxy-.-[methyl] butanoic acid (LMA: liquid form).
Both powder and liquid forms consist of an L-isomer and
a D-isomer at a ratio of +: +. In the metabolic pathway of
poultry, 1*+** of the D-isomer of DLM or LMA is
converted to L-isomer (Baker and Boebel, +32*; Noll et
al., +32.; Baker, +33.; Hasegawa et al., ,**/).
Comparative Bioe$cacy of DLM and LMA
The average relative bioavailability of LMA compared
with DLM in various animal species is approximately 0/
3* (Reid et al., +32,; van Weerden and Schutte, +32.;
Scott, +321; Esteve-Garcia and Austic, +33-b; Harms and
Russell, +33.; Hoehler and Hooge, ,**-). The nutritional
e#ectiveness of LMA relative to that of DLM and the
reasons for the di#erence have been widely investigated
(Esteve-Garcia and Austic, +33-a; Huyghebaert, +33-;
Rostagno and Barbosa, +33/; Lemme et al., ,**,). Nevertheless, the exact metabolic processes that result in the
bioavailability di#erence remain unclear (Liu et al., ,**.).
Recently, data from .0 dose-response experiments extracted from a total of ,1 peer-reviewed papers were
analyzed, and the meta-analysis showed that the biological
e$ciencies of LMA were 2+ and 13 of the values for
DLM, on an equimolar basis, for weight gain and feed
conversion ratio, respectively (Sauer et al., ,**2). Regression analysis also revealed that LMA was 02 (weight
gain), 01 (feed conversion), 0, (carcass yield) and
0. (breast meat yield) as e$cacious as pure DLM on an
as-fed basis, meanwhile LMA was 1, (weight gain),
/+ (feed conversion), .2 (carcass yield), and 0*
(breast yield) as e$cacious as DLM on a weight-forweight basis (Lemme et al., ,**,). However, Bunchasak
et al. (,**0) concluded that LMA is an acceptable source

of methionine (22 bioe$cacy) for broiler chicks fed

corn-soybean meal diets from day-old to market weight, if
maximal growth performance is used as a response criterion concerning the commercial methionine requirement
recommendation (total sulfur amino acids (TSAA) requirement+.*- and *.3- during starter and grower
periods, respectively) (Table +). Similarly, V
azquezAn

on et al. (,**0) compared the gain-response curve to

dietary levels of LMA and DLM and concluded that the
two methionine sources have di#erent dose-response
forms, LMA outperforms DLM at commercial levels,
whereas DLM outperforms LMA at decient levels.
Therefore, it can be concluded that the bioe$cacy of
LMA would be 22 of DLM (equimolar basis) if the
commercial level of TSAA is set in the formulation of a
diet.
Absorption and Transportation
Normally, amino acids, including methionine, are
mainly absorbed through the small intestine. Two to three
decades ago, some reports state that L-methionine is transported by a single Na-dependent mechanism (Lerner
et al., +31/; Knight and Dibner, +32.; Brachet and
Puigserver, +323). Recently, however, Soriano-Garc
a et
al. (+332) stated that the absorption of dipolar amino
acids such as L-methionine by the small intestine (brushborder membrane vesicles) in chicken is mediated by
multiple pathways. They reported that L-methionine is
transported by systems specic to neutral amino acids
(L-like: Na-independent, and B-like: Na-dependent)
and by systems that also transport cationic amino acids
(ym-like: high Vmax for L-methionine transport in the
absence of Na and b*, -like: interaction with cationic
amino acids) (Soriano-Garc
a et al., +332, +333). Using
molecular biotechnology to determine the transportation
also indicated multiply sized poly (A) RNA that encodes
proteins capable of mediated transport of L-methionine
and/or LMA present in broiler intestinal mucosa (Pan et
al., ,**,). Thus, the absorption and transportation of
methionine are complicated processes.
Regarding age, Noy and Sklan (+330) reported that

The e#ects of dietary treatments on growth performance of male broiler chickens from
*0 weeks of age

Table +.

Relative bioe$cacy of LMA to DLM

Initial weight (g)

Final weight (g)
Weight gain (g)
Feed intake (g/day)
Feed conversion ratio
Mortality ()
Uniformity ()

Basal diet

DLM

-240
,5-10a
,5--2a
.5*-*a
+41,a
*4**
224*2a

-240
,51,/b
,5021b
.5,11b
+4/3b
*4**
3*4..ab

10

2*

2.

22

-240
,51,+b
,502,b
.5,02b
+4/3b
+4-3*4+3ab

-240
,5123b
,51/+b
.5.3+b
+40-b
*401
3,4-+b

-240
,51-/b
,5031b
.5-+3b
+40*b
+4-3*40/b

-240
,52*.b
,510/b
.5.32b
+40-b
-4-3+40+b

a, b
: Means within a row with no common superscripts di#er signicantly (P*.*/).
Adapted from Bunchasak et al. (,**0).

Bunchasak: Methionine for Poultry Production

methionine uptake capacity increased in both the duodenum and jejunum between hatching and 1 days of age, and
remained constant between 1 and +. days of age, so they
postulated that from 1 days of age feed intake may be the
major factor controlling nutrient uptake in chicks. Excess
methionine supplementation seems to reduce the potential
of the uptake of methionine itself and other nutrients.
Soriano-Garc
a et al. (+333) reported that excess methionine supplementation downregulates specic transport
mechanisms of the small intestine involved in the apical
L-methionine transport. Moreover, L-lysine uptake is
strongly inhibited by excess methionine supplementation
(Torras-Llort et al., +330, +332), and excess SAA a#ects
organic bone matrix metabolism and may play a role in the
etiology of tibial dyschondroplasia (Frankel, +33/).
By focusing on di#erences between methionine sources,
di#erent multiple transport systems appear to be involved
in transporting both DLM and LMA. Knight and Dibner
(+32.) reported that L-methionine absorption may be
accomplished by both concentration- and energydependent processes, while the absorption of LMA is
concentration-dependent. Pan et al. (,**,), Dibner et al.
(+322), Knight and Dibner (+32.) and Brachet and
Puigserver (+321) also suggested that mechanisms of
methionine absorption include Na-dependent transport,
Na-independent transport and/or di#usion, while the
mechanisms suggested for LMA absorption include Naindependent but H-dependent transport and/or di#usion.
Therefore, di#erences in the mechanism of transportation
between the two methionine sources may lead to a di#erence in the amount of their transportation.
Although the ability to use LMA as a source of
methionine would not be limited by absorption (Knight
and Dibner, +32.), it has been reported that the small
intestine has similar capacities to absorb LMA and Lmethionine (Knight and Dibner, +32.; Han et al., +33*).
Moreover, the lower intestinal absorption of LMA relative
to DLM (Esteve-Garcia and Austic, +33-a; Lingens and
Molnar, +330; Maenz and Engele-Schaan, +330) has also
been reported. The reasons given for this are: +) Lmethionine has a higher a$nity for its transporter and a
higher maximal velocity of transport than LMA, and
L-methionine is removed more quickly from the intestinal
lumen than LMA and therefore has less exposure to
intestinal bacteria resulting in decreased methionine
uptake (Drew et al., ,**-), and ,) high amounts of
derivative products (cysteine, glutathionine or taurine)
may be produced from LMA rather than DLM (Mart
nVenegas et al., ,**0).
Interrelationship between
Methionine and Other Nutrients
Dietary Cyst(e)ine and Sulfur
Among essential amino acids, methionine seems to have
many interrelationships with other nutrients because
many metabolic pathways and other nutrients (cystine,
choline, betain, vitamin B0, vitamin B+, and folate) involve

171

methionine (Bunchasak, ,**2). The molar e$cacies of

methionine, +/, cystine and cysteine were the same
(Graber and Baker, +31+). Poultry require both methionine and cysteine for protein synthesis, so the TSAA
requirement should be taken into account. In avian
species, it is generally accepted that around ./ to /* of
TSAA can be supplied by cysteine. However, cystine supplementation has a negative impact on voluntary feed
intake when the diet is markedly decient in TSAA (i.e.
when greater than /* of TSAA intake is provided by
cystine) (Dilger and Baker, ,**1). Moreover, when the
TSAA requirement is expressed as a percentage of the
diet, the need for a methioninecystine combination is
less than that for methionine alone (Graber and Baker,
+31+). Compared to DLM, a +: + mixture of DLM and
L-cysteine was only 2+ or 20 as e#ective in supporting
growth or food conversion, respectively (Huyghebaert
and Pack, +330).
Since TSAA also act as a sulfur donor, sulfur supplementation inuences the sparing e#ect between methionine and cysteine. It seems that when sulfur sources are
added to the diet, the sparing e#ect between cysteine and
methionine or the TSAA requirement is reduced. Sasse
and Baker (,***) found that when TSAA were at or near
adequacy, the optimal percentage from cystine was .2..
in the presence of dietary K,SO. and /,.1 in its absence,
whereas when TSAA were set at a decient level, optimal
performance occurred when cystine furnished .+.- and
.2., of the TSAA need, respectively, in the presence
and absence of K,SO., respectively. Using practical broiler nisher diets, three trials were carried out to determine
to what extent synthetic methionine can be replaced by
sodium sulphate. The results revealed that weight gain
and the feed conversion ratio both increased with incremental increases in sodium sulphate in diets containing
sub-optimal concentrations of TSAA (Plavnik and
Bornstein, +312).
Methyl Donors
Methionine, betaine and choline are methyl donors that
play important roles in methylation reactions, their interrelationship has been illustrated by Pillai et al. (,**0).
Methionine has been added to overcome growth depression even when caused by dietary tannic acid and mild
arginine toxicity via the property of the methyl donor
(Kim et al., ,**0). It seems that the e#ect of methionine
is unique and that other methyl donors could not substitute for methionine, since many researchers have reported
that betaine and choline can only slightly or not substitute
for methionine in broilers or laying hens fed diets marginally decient in TSAA (McDevitt et al., ,***; Molitoris
and Baker, +310; Baker et al., +32-; Pourreza and Smith,
+322; Rostagno and Pack, +330; Schutte et al., +331).
For slow-growing chicks (+..0 g/day), however, Attia et
al. (,**/) reported that the methionine level can be
decreased to *.-1 or even to *.-, of the diet when
*.*1 betaine is added. This replacement potential may
be the result of a low methionine requirement due to the

J. Poult. Sci., .0 (-)

172

slow growth rate. Moreover, increasing carcass fat by

manipulating the percentage dietary protein level or the
amino acid balance (methionine deciency) does not inuence betaines activity as a lipotropic agent (Garcia Neto
et al., ,***).
Protein Level in the Diet
The optimal level of methionine in the diet seems to
depend on the protein concentration in the diet. Vieira et
al. (,**.) conducted an experiment with +,..* male Cobb
/** and +,..* male Ross -*2 broilers (+. to -/ days of
age) and suggested that the optimum dietary TSAA level
depends on the dietary protein level. The TSAA requirement does not change with age when it is expressed in
terms of dietary protein (Bornstein and Lipstein, +300).
In addition, Mendonca and Jensen (+323) and
Huyghebaert and Pack (+330) reported that a broiler
chicks requirement for TSAA increased with increasing
dietary protein concentrations ranging from +3.1 to
,/.3. Therefore, several investigators have suggested
that the methionine concentration in chick diets should be
around ,./. of the protein concentration (Boomgaardt
and Baker, +31-; Mendonca and Jensen, +323; Morris et
al., +33,).
Although an increase in the dietary methionine requirement is often found with elevated protein concentrations,
the capacity to use methionine for protein gain is also
reduced (Fatufe and Rodehutscord, ,**/). Sterling et al.
(,**0) intensively reviewed the ratio of protein: amino
acids, and found that the amino acid requirements expressed as a percentage of diet tended to decline as protein
content increased. In laying hens, when the ratio of
protein: methionine was kept constant, methionine supplementation to a high protein level (+2 CP) depressed
egg production, while supplementation to lower protein
levels (+. and +0 CP) improved the production performance (Poeikhampa, ,**.). These results may suggest
that increasing the cysteine content by increasing dietary
protein concentration reduces the methionine requirement. Thus, the positive e#ect of keeping a constant
protein: methionine ratio occurs when the appropriate
range of protein is provided.
Methionine and Heat Stress
High environmental temperature decreases the feed
intake to maintain homeothermy (Gonzalez-Esquerra and
Leeson, ,**0a), and the subsequent live weight gain,
digestibility, egg production, egg quality and feed e$ciency are poor (Austic, +32/; Bunchasak and Silapasorn,
,**/). Diets with an amino acid imbalance or methionine
deciency normally increase heat production (Sekiz et al.,
+31/), and induce a more negative e#ect of heat stress
when the environmental temperature is high (Bunchasak
and Silapasorn, ,**/). As balancing the amino acid
composition in the diet with methionine supplementation
improves production performance through pathways of
polyamine metabolism (Gonzalez-Esquerra and Leeson,
,**0a), glutathionine (derived from methionine) may

reduce damage from oxidative stress. So, the TSAA

requirement would be higher under hyper-thermoneutral
conditions than under thermoneutral conditions. Silva et
al. (,**0) who conducted two trials to determine the
TSAA requirement for Ross female broilers from + to ,+
and ,, to ., days of age reported that raising broiler
chicks at high temperature requires higher TSAA consumption to achieve optimal growth performance.
It is well established that dietary protein produces a
high heat increment. Austic (+32/), Waldroup et al.
(+310) and Waldroup (+32,), therefore, recommend a
reduction in dietary protein content with suitable supplementation by essential amino acids to alleviate the
negative e#ects of heat stress. However, the reduction of
limiting amino acid or protein content in the diet
negatively a#ects production performance (Morris and
Gous, +322; Bunchasak and Silapasorn, ,**/). For example, the weight of eggs produced by hens more than -2
weeks of age was reduced by limiting methionine intake to
,1* mg per hen daily, compared with -** mg methionine
per hen daily (Peterson et al., +32-). On the other hand,
methionine supplementation to low-protein diets improves
production performance (Sell and Rogler, +32-; Summers
et al., +33+; Harms and Russell, +33-; Chung et al., +332;
Ravikiran and Devegowda, +332). Therefore, reducing
the dietary protein concentration by methionine supplementation reduces the negative e#ects of heat stress.
Gonzalez-Esquerra and Leeson (,**0b) reported that
Arg: Lys, methionine source and duration of exposure to
heat stress a#ected protein utilization in hyperthermic
birds. Bunchasak and Silapasorn (,**/) found that
methionine intake of .-3.3- mg/hen/day (+. CP;
*... methionine) improved hen-day egg production and
egg weight to the level of a control group (+0 CP;
*.-2 Met), which had a methionine intake of -1,.3.
mg/hen/day (Table ,). Moreover, adding methionine to
a low-protein diet reduced the mortality rate of hens under
heat stress when compared to a positive control group
(+0 CP; *.-2 methionine) or a negative control (+.
CP; *.,0 methionine). However, Amaefule et al.
(,**.) used old Bovan Nera layers that had been in lay
for .* weeks to evaluate the e#ect of methionine, lysine
and/or vitamin C supplementation on egg production as
well as on external and internal egg quality characteristics
of layers in a humid tropical zone, and reported that none
of these supplements had any benet to the layer hens.
Thus, age and production conditions may be factors to
consider when adding methionine in order to reduce the
negative e#ect of heat stress.
Rostagno and Barbosa (+33/) compared the absorption
rate of DLM and LMA during heat stress and found that
net absorption of LMA was signicantly lower than that
of DLM (broiler chicks ,+ to ., days of age). Recently,
Mitchell and Lemme (,**2) reported that at high temperature (-/), the length and weight of the small intestine
decreased while the absorption rate of both methionine
sources increased compared to that under thermoneutral

Bunchasak: Methionine for Poultry Production

173

E#ects of additional methionine in low-protein diet on production performance of laying hens from ,. to .. weeks of age under heat stress (-/)

Table ,.

Item

Feed intake (g/day)

Protein intake (g/day)
Met intake (mg/day)
Egg production ()
Egg weight (g)
Egg mass (g/hen/day)
FCR (g feed /g egg)
Mortality ()

Control
(+0 CP)
*4-2 Met
324+.a
+/41*a
-1,43.b
1/40*a
/.43,a
.,4*/a
,4--c
24,,b

Low-CP diet (+. CP)

*4..
Met
33432a
+.4**b
.-343-a
1.410a
/.42,a
.*432b
,4.-b
/421c

*4-2
Met
3341-a
+-430b
-12430b
1,4/+b
//4,,a
.*4.+b
,4.1b
/423c

*4-*
Met
324,+a
+-41/b
,3.40.c
034+0b
/.4.-a
-140.c
,40*a
04*1b

*4,0
Met
2142/b
+,4-*c
,,24.*d
0/4,.c
/*42*b
--4-.d
,40-a
++41/a

ad

: Means within a row with no common superscripts di#er signicantly (P*.*/).

Adapted from Bunchasak and Silapasorn (,**/).

conditions (,,). They also suggested a higher absorption velocity for DLM than that for LMA. In another
study, however, when broilers (-0 weeks of age) were
kept at a constant -* or at diurnally cycling temperatures of ,/-/, the di#erences in response to DLM and
LMA were small (Balnave and Oliva, +33*). DLM and
LMA addition to vegetable-based diets in comparable
molar terms also promoted similar performance in broilers
under chronic heat stress conditions (Ribeiro et al., ,**/).
In contrast, Knight et al. (+33.) indicated that the
capacity for DLM uptake into intestinal epithelial cells
and DLM utilization were reduced in heat-stressed birds,
whereas the LMA uptake and utilization were not
reduced. This is because energy-independent uptake is
increased during heat stress, but not enough to compensate for the decrease in carrier-specic energy- and
sodium-dependent uptake during heat stress (Dibner et
al., +33,). Additionally, it has been reported that under
acute heat stress conditions the choice of a supplemental
source of methionine activity should take into account the
dietary Arg: Lys ratio (Chen et al., ,**-; Balnave et al.,
+333). Therefore, the best choice of a methionine source
under heat stress is still unclear.
E#ect of Methionine on the Immune System
Grimble (,**0) has intensively reviewed the e#ects of
TSAAs and their major metabolites on health and disease
in humans. Their e#ect can be divided into two routes:
the rst is a su$cient metabolic supply of TSAA from the
diet and tissue protein breakdown that supports the synthesis of myriad proteins and peptides involved in normal
functioning of the immune system, and the second is three
major products of TSAA (glutathione, homocysteine and
taurine) that inuence inammatory aspects of the
immune response (Grimble, ,**0).
There are some reports that high methionine supplementation promotes good health for poultry. For example, the supplementation improved leukocyte migration
inhibition, cellular immune response and humoral immune

response (Swain and Johri, ,***; Attia et al., ,**/), increased blood serum total protein, albumin, globulin and
antibody response to Newcastle disease virus, and
decreased serum aspartate aminotransferase and alanin
aminotransferase (Attia et al., ,**/), increased total antibody, IgG, and response to the mitogen phytohemagglutinin (PHA), which might be related to T-cell help
(Tsiagbe et al., +321). So, methionine deciency results in
decreases in both the humoral and nonspecic immunocompetence of broiler chickens (Zhang and Guo, ,**2).
In long-term depression of dietary protein, our study
suggested that adequate dietary methionine is necessary
for sustaining normal immunocompetence and achieving
maximum production performance (Poosuwan et al.,
,**3). Additionally, Takahashi et al. (+331) suggested
that dietary cysteine also has an impact on the immune
and inammatory responses. However, on an equimolar
basis (+0.2 mmol/kg diet), cysteine is about 2. and 1*
as e$cacious as methionine in IgG and PHA stimulation,
respectively (Tsiagbe et al., +321).
Moreover, methionine has been added to overcome
growth depression even when the depression is caused by
dietary tannic acid and mild arginine toxicity via the
property of the methyl donor (Kim et al., ,**0).
Additionally, TSAA supplementation of diets containing
aatoxin improved performance in chickens (Veltmann et
al., +32+, +32-), while this response seems to be less in
weanling pigs (van Heugten et al., +33.). Both forms of
methionine (DLM and LMA) reduced intestinal populations of Clostridium perfringens in broiler chickens when
used in relatively high concentrations, and may reduce the
risk of necrotic enteritis (Dahiya et al., ,**1). Recently,
Xie et al. (,**1) reported that the toxicity of LMA was
low relative to that of DLM. Matsushita et al. (,**1)
suggested that marginal excess supplementation of diet
with LMA in place of DLM improved meat production in
female broiler chicks, and that dietary LMA has the
potential to alleviate certain stress responses. These
results suggest that LMA is converted into glutathione,

J. Poult. Sci., .0 (-)

174

homocysteine and taurine more easily than DLM, since

the cysteine and taurine content in enterocytes is higher
when LMA is used as the methionine source (Mart
nVenegas et al., ,**0).
Methionine Requirement
Broiler Chickens
Feed consumption is mainly controlled by dietary
energy. Summers et al. (+33,) reported that the level and
balance of essential amino acids (EAA) also signicantly
a#ected feed intake, consequent weight gain and carcass
composition. Broiler chicks appear to react to amino acid
deciencies within a short period (hours) by adjusting
their feed intake and/or selection and these responses are
inuenced by age and prior experience (Picard et al.,
+33-). However, Bunchasak et al. (+330, +332) and
Bunchasak and Keawarun (,**0) found that methionine
deciencies depressed the feed intake of broiler chicks due
to amino acid imbalances. It can be assumed that, under
amino acid imbalances, chicks lose the potential to adjust
feed intake to satisfy their amino acid requirements; the
main positive e#ect of methionine supplementation may
come from its improvement of feed intake via the amino
acid balance.
Schutte and Pack (+33/) designed two experiments to
evaluate the e#ect of methionine and TSAA in a cornsoybean diet on broiler performance and carcass yield.
They found that the TSAA requirement was higher for
obtaining maximum feed utilization and breast meat yield
than for obtaining maximum weight gain. Atencio et al.
(,**.) reported that TSAA levels had positive linear and
quadratic e#ects on body weight gain and feed conversion,
Table -.

respectively, at + to ,* days of age. Using the broken line

model, they estimated that the requirement of broiler
chicks as digestible TSAA or the percentage total in the
diet during + to ,* days of age was *.2*2 or *.3*1 total,
while the ratio of digestible TSAA: lysine for body weight
gain and for the feed conversion ratio (FCR) was 1*
during ,.-2 days of age, those values were *.101 or
*.2.. total and 1,, respectively, and during ../0
days of age they were *.002 or *.1-2 total and 1+,
respectively. Under European price conditions, based on
the feed conversion ratio, the dietary level of *.3/
TSAA was found to be most protable for broilers grown
to +.1 kg, while in broilers grown to ,., kg, the most
protable TSAA level would be *.2/ (Schutte and Pack,
+33/). Commercially, the recommended methionine requirement is always higher than the NRC (+33.) recommendation. The methionine requirement expressed as a
percentage of diet declines when chicks are grown up,
while the patterns of methionine requirement (relative to
lysine) change little (but tend to increase) (Table -).
Genetic diversity also inuences the methionine utilization of chickens. Consequently di#erent strains require
di#erent amounts of methionine (see Table -). Geraert et
al. (+321) observed that the genetically fat-type chicken
had lower plasma concentrations of most glucogenic
amino acids and higher levels of branched-chain and
sulfur-containing amino acids than lean-type chicken.
Leclercq et al. (+33-) developed two lines of chickens,
lean (LL) and fat (FL) male chickens, and fed them ve
diets di#ering by TSAA content (/.., /.2, 0.,, 0.0 and 1.*
g/kg) from ,2 to .1 days of age. They also found that
lean chickens required a higher dietary concentration of

Broiler nutrition recommendations according to the nutrition guideline of strains

Nutrition recommendation ()

Strain

Stage

TSAA

Relative to lysine

Methionine
TSAA

Methionine

*4.1
*4.,
*4-2

1.
10
12

-2
-2
-3

*4/0
*4/*4.2
*4..

*4/*
*4.2
*4.*4.*

1.
1/
12
12

-2
.*
.+
.+

*420
*41/
*403
*402

*4/*4.0
*4.,
*4.,

*4.0
*4.+
*4-1
*4-1

1+
1,
110

-3
-3
-3
.+

*4/*
*4-2
*4-,

2,
1,
1+

.0
-2
-2

Total

Digest

Total

Digest

Starter
Grower
Finisher

+4*1
*43/
*40*

*43.
*42.
*410

*4/+
*4./
*4.+

Starter
Grower
Finisher+
Finisher,

*432
*430
*422
*42*

*420
*42.
*411
*41

ArborAcres

Starter
Grower
Finisher+
Finisher,

*431
*42/
*412
*411

NRC (+33.).

Starter
Grower
Finisher

*43*
*41,
*40*

Ross -*2+

Cobb

Starter
Starter
Starter
.
Starter
,

(*+*
(*+*
(*+.
(*,+

days),
days),
days),
days),

grower
grower
grower
grower

(++,.
(++,,
(+/,2
(,,.,

days) and nisher (,/-slaughter).

days), nisher+ (,-., days) and nisher, (.,-slaughter).
days), nisher+ (,3-0 days) and nisher, (-1-slaughter).
days) and nisher (.-/0 days).

Bunchasak: Methionine for Poultry Production

TSAA and used TSAA more e$ciently than FL. Moreover, LL chickens require diets more concentrated in nonessential amino acids than fat chickens (Alleman et al.,
,***). However, the methionine requirement of Na/na
(Naked neck gene) chickens did not di#er from that of
their normally feathered counterparts under either spring
or summer ambient temperature conditions (Yalcin et al.,
+333).
Many factors such as genetic diversity, environmental
conditions, nutrients and stress involve body fat deposition. Summers et al. (+33,) reported that the level and
balance of EAA have a signicant e#ect on feed intake,
thereby inuencing weight gain and carcass composition.
Generally, larger chickens have more breast meat and a
heavier abdominal fat pad. Increasing dietary methionine
increases the mass of breast meat but reduces the size of
the abdominal fat pad due to a good balance of amino
acids (Wallis, +333; Bunchasak et al., +330, +332).
Additionally, Zhan et al. (,**0) reported that methionine
supplementation signicantly increased breast muscle
yield and decreased abdominal fat content. They found
that supplementation with methionine signicantly increased the contents of creatine and free carnitine in the
liver, the activity of hormone-sensitive lipase in abdominal
fat and the concentration of free fatty acid in serum,
whereas uric acid concentration in serum was signicantly
decreased. Therefore, the decrease in abdominal fat may
be due to increased carnitine synthesis in the liver and
hormone-sensitive lipase activity in abdominal fat.
Laying Hens
Unlike broiler chickens, the methionine requirement of
laying hens should be expressed as mg/day. For white-egg
laying hens, a requirement of approximately 11/ to 2** mg
TSAA/hen/days of which about -3* to ..* mg was
methionine, was found for a maximum of 2* to 2- eggs/
+** hen/days (Schutte and van Weerden, +312). NRC
(+33.) reported that the white-egg hens require -** and
/2* mg of methionine and TSAA per hen daily, respectively, while the methionine and TSAA requirement for the
brown egg type of laying hens is --* and 0./ mg/hen/day,
respectively. However, several investigators reported a
higher requirement of methionine for maximal egg production. Bunchasak and Silapasorn (,**/), Cao et al.
(+33,), Schutte et al. (+33.) and Novak et al. (,**.)
found that the methionine and TSAA requirements were
around .,...* and 1.*2++ mg per hen daily, respectively. Thus, white-egg laying hens require lower TSAA than
brown-egg laying hens, and commercial laying hens require higher TSAA than the NRC (+33.) recommendation.
Consumption above .+- mg/day methionine resulted in
signicantly increased albumen total solids and protein,
and yolk protein was signicantly increased at /*1 and //0
mg/day methionine compared to .+- mg/day methionine
(Shafer et al., +332). The methionine and TSAA requirements were greater for the middle and nal quarters of
production than for the initial quarter (Waldroup and

175

Hellwig, +33/).
The peak daily requirements for
methionine were -2., -2* and .*, mg/day for egg production, egg weight and egg mass, respectively (Waldroup
and Hellwig, +33/). These data indicate that the requirement for maximum egg production is less than that for
maximum feed utilization (Schutte and van Weerden,
+312; Schutte, et al., +32-) and the requirement for egg
quality is higher than that for egg production and feed
utilization. A study of the methionine requirement of
Isa-brown laying hens conducted in Australia found that a
dietary methionine concentration of ,2- mg/kg or *.,2-
(mean daily methionine intake was -1* mg and .-- mg for
hens housed in single- and multiple-hen (/ hens/cage)
cages, respectively) is su$cient to satisfy the methionine
requirements (Poultry Research Foundation, ,***).
These uncertain results may be due to a number of
factors such as environmental conditions, the management system and dietary protein or energy levels. Therefore, Waldroup and Hellwig (+33/) suggested that adjustments should be made in dietary amino acid levels to
compensate for changes in daily feed intake as inuenced
by environmental changes, feather covering or other
factors in order to maintain a constant amino acid intake,
but adjustment based on age or stage of production is not
justied. However, based on a review of the literatures, it
seems that methionine intake should be higher than .,*
mg/day to maximize the quantity and quality of egg
production.
Conclusions
The use of synthetic methionine is very important for
poultry production. The level of supplementation should
be carefully considered due to a variety of factors. The
amount of methionine required to support the immune
system seems to be high because it has to be used not only
for protein synthesis but also to produce some antioxidants. The di#erence between DLM and LMA in absorption and utilization abilities under heat stress is still unclear because the absorption system is very complicated.
However, in normal situations, LMA can be used as a
source of methionine.
Acknowledgments
The author gratefully acknowledges and appreciates the
support of Dr. Atsuro Matsuda, Sumitomo Chemical Co.,
Ltd., Japan.
References
Aldrich G. DL-methionine: several vital functions. Available
Source: http: / / www. petfoodindustry. com / ViewArticle.
aspx?id+2/,0, ,**1, October ,3.
Alleman F, Michel J, Chagneau AM and Leclercq B. The e#ects
of dietary protein independent of essential amino acids on
growth and body composition in genetically lean and fat
chickens. British Poultry Science, .+: ,+.,+2. ,***.
Amaefule KU, Ojewola GS and Uchegbu EC. The e#ect of
methionine, lysine and/or vitamin C (ascorbic acid) supple-

176

J. Poult. Sci., .0 (-)

mentation on egg production and egg quality characteristics of layers in the humid tropics. Livestock Research for
Rural Development, ,**.. Available Source: http://www.
lrrd.org/lrrd+0/3/cont+0*3.htm
Atencio A, Fernando L, Albino T, Rostagno HS and Vieites FM.
Methionine cystine requirement of male broilers from +
to ,*, ,. to -2 and .. to /0 days of age. Revista Brasileira
de Zootecnia, --: ++/,++00. ,**..
Attia YA, Hassan RA, Shehatta MH and Abd El-Hady Slawa B.
Growth, carcass quality and serum constituents of slow
growing chicks as a#ected by betaine addition to diets
containing ,. Di#erent levels of methionine. International
Journal of Poultry Science, .: 2/020/. ,**/.
Austic RE. Feeding poultry in hot and cold climates. In: Stress
Physiology in Livestock (Youssef MK ed). pp. +,-+-0.
CRC Press. Boca Raton FL. +32/.
Balnave D and Oliva A. Responses of nishing broilers at high
temperatures to dietary methionine source and supplementation levels. Australian Journal of Agricultural Research, .+:
//1/0.. +33*.
Balnave D, Hayat J and Brake J. Dietary arginine: lysine ratio
and methionine activity at elevated environmental temperatures. Journal of Applied Poultry Research, 2: +3. +333.
Baker DH and Boebel KP. Utilization of the D- and L-isomers of
methionine and methionine hydroxy analogue as determined
by chick bioassay. Journal of Nutrition, ++*: 3/330.. +32*.
Baker DH, Halpin KM, Czarnecki GL and Parsons CM. The
choline-methionine interrelationship for growth of the
chick. Poultry Science, 0,: +--+-1. +32-.
Baker DH. Utilization of precursors for L-amino acids. In:
Amino acids in farm animal nutrition (DMello JPF ed).
pp. -10,. CAB International. Wallingford, UK. +33..
Boomgaardt J and Baker DH. E#ect of dietary energy concentration on sulfur amino acid requirements and body composition of young chicks. Journal of Animal Science, -0: -*1
-++. +31-.
Bornstein S and Lipstein B. Methionine supplementation of
practical broiler rations: III. The value of added methionine
in broiler nisher rations. British Poultry Science, 1: ,1-
,2.. +300.
Brachet P and Puigserver A. Transport of methionine hydroxyl
analog across the brush border membrane of rat jejunum.
Journal of Nutrition, ++1: +,.++,.0. +321.
Brachet P and Puigserver A. Na-independent and nonstereospecic transport of ,-hydroxy-.-methylthiobutanoic
acid by brush border membrane vesicles from chick small
intestine. Comparative Biochemistry and Physiology, 3.B:
+/1+0-. +323.
Bunchasak C, Tanaka K, Ohtani S and Collado CM. E#ect of
metcys supplementation to a low-protein diet on the
growth performance and fat accumulation of broiler chicks
at starter period. Animal Science and Technology (Japan),
01: 3/0300. +330.
Bunchasak C, Tanaka K and Ohtani S. E#ect of supplementing
nonessential amino acids on growth performance and fat
accumulation in broiler chicks fed a diet supplemented with
MetCys. Japanese Poultry Science, -/: +2,+22. +332.
Bunchasak C and Silapasorn T. E#ects of adding methionine in
low-protein diet on production performance, reproductive
organs and chemical liver composition of laying hens under
tropical conditions. International Journal of Poultry Science, .: -*+-*2. ,**/.

Bunchasak C and Keawarun N. E#ect of methionine hydroxy

analog-free acid on growth performance and chemical composition of liver of broiler chicks fed a corn-soybean based
diet from * to 0 weeks of age. Animal Science Journal, 11:
3/+*,. ,**0.
Bunchasak C, Sooksridang T and Chaiyapit R. E#ect of adding
methionine hydroxyl analogue as methionine source at the
commercial requirement recommendation on production
performance and evidence of ascites syndrome of male
broiler chicks fed corn-soybean based. International Journal
of Poultry Science, /: 1..1/,. ,**0.
Bunchasak C. Using synthetic amino acids in broiler chicken
diet. In: Proceedings of Satellite Symposium, International
Symposium on the Future & Practical Usage of Amino
Acids for Animal Production. XIII AAAP Animal Science
Congress. pp. 1+2. ,**2.
Cao Z, Jevne CJ and Coon CN. The methionine requirement of
laying hens as a#ected by dietary protein levels. Poultry
Science, 1+(supplement+): -3 (Abstract). +33,.
Chadd SA, Davies WP and Jason MK. Practical production of
protein for food animals. In: Proceedings of Protein Sources
for the Animal Feed Industry, Expert Consultation and
Workshop Bangkok, FAO Animal Production and Health.
,**,. Available Source: http://www.fao.org/docrep/**1/y
/*+3e/y/*+3e*1.htm#bm*1
Chavez C, Coufal CD, Lacey RE and Carey JB. The impact of
methionine source on poultry fecal matter odor volatiles.
Poultry Science, 2-: -/3-0.. ,**..
Chen J, Hayat J, Huang B, Balnaved D and Brake J. Responses
of broilers at moderate or high temperatures to dietary
arginine: lysine ratio and source of supplemental methionine
activity. Australian Journal of Agricultural Research, /.:
+11+2+. ,**-.
Chung HJ, Chung-Yi L and Wen-Shyg Chiou P. E#ects of
ambient temperature and methionine supplementation of a
low protein diet on the performance of laying hens. Animal
Feed Science and Technology, 1.: ,23,33. +332.
Dahiya JP, Hoehler D, Van Kessel AG and Drew MD. E#ect of
di#erent dietary methionine sources on intestinal microbial
populations in broiler chickens. Poultry Science, 20: ,-/2
,-00. ,**1.
Dibner JJ, Knight CD, Swick RA and Ivey FJ. Absorption of +.
C-,-hydroxy-.-(methylthio) butanoic acid (Alimet) from
the hindgut of the broiler chick. Poultry Science, 01: +-+.
+-,+. +322.
Dibner JJ, Atwell CA and Ivey FJ. E#ect of heat stress on ,hydroxy-.-(methylthio) butanoic acid and DL-methionine
absorption measured in vitro. Poultry Science, 1+: +3**
+3+*. +33,.
Dilger RN and Baker DH. DL-Methionine is as e$cacious as
L-methionine, but modest L-cystine excesses are anorexigenic in sulfur amino acid-decient puried and practicaltype diets fed to chicks. Poultry Science, 20: ,-01,-1..
,**1.
Drew MD, Van Kessel AG and Maenz DD. Absorption of
methionine and ,-hydroxy-.-methylthiobutoanic acid in
conventional and germ-free chickens Poultry Science, 2,:
++.3++/-. ,**-.
Esteve-Garcia E and Austic RE. Intestinal absorption and renal
excretion of dietary methionine sources by the growing
chicken. Journal of Nutritional Biochemistry, .: /10/21.
+33-a.

Bunchasak: Methionine for Poultry Production

Esteve-Garcia E and Austic RE. Performance, breast meat yield

and abdominal fat deposition of male broiler chickens fed
diets supplemented with DL-methionine or DL-methionine
hydroxy analogue free acid. British Poultry Science, -.:
-/+-/3. +33-b.
Fatufe AA and Rodehutscord M. Growth, body composition,
and marginal e$ciency of methionine utilization are a#ected
by nonessential amino acid nitrogen supplementation in
male broiler chicken. Poultry Science, 2.: +/2.+/3,. ,**/.
Frankel TL. Sulfate incorporation into organic bone matrix of
the tibiotarsus of broiler chicks is reduced by excess dietary
methionine. Poultry Science, 1.: /+*/+0. +33/.
Garcia Neto M, Pesti GM and Bakalli RI. Inuence of dietary
protein level on the broiler chickens response to methionine
and betaine supplements. Poultry Science, 13: +.12+.2..
,***.
Geraert PA, Leclercq B and Larbier M. E#ects of dietary
glucogenic amino acid supplementation on growth performance, body composition and plasma free amino acid levels in
genetically lean and fat chickens. Reproduction, Nutrition
and Development, ,1: +*.++*/+. +321.
Gonzalez-Esquerra R and Leeson S. Physiological and metabolic
responses of broilers to heat stress - implications for protein
and amino acid nutrition. Worlds Poultry Science Journal,
0,: ,2,,3/. ,**0a.
Gonzalez-Esquerra R and Lesson S. E#ect of arginine: lysine
ratios and source of methionine on growth and body protein
accretion in acutely and chronically heat-stressed broilers.
Poultry Science, 2/: +/3.+0*,. ,**0b.
Graber G and Baker DH. Sulfur amino acid nutrition of the
growing chick: quantitative aspects concerning the e$cacy
of dietary methionine, cysteine and cystine. Journal of
Animal Science, --: +**/+*++. +31+.
Grimble RF. The E#ects of sulfur amino acid intake on immune
function in humans. Journal of Nutrition, +-0: +00*S+00/S.
,**0.
Han Y, Castanon F, Parsons CM and Baker DH. Absorption and
bioavailability of DL-methionine hydroxy analog compared
to DL-methionine. Poultry Science, 03: ,2+,21. +33*.
Han Y and Baker DH. E#ects of excess methionine or lysine for
broilers fed a corn-soybean meal diet. Poultry Science, 1,:
+*1*+*1.. +33-.
Harms RH and Russell GB. Optimizing egg mass with amino
acid supplementation of a low-protein diet. Poultry Science,
1,: +23,+230. +33-.
Harms RH and Russell GB. A comparison of the bioavailability
of DL-methionine and methionine hydroxy analogue acid
for the commercial laying hen. Journal of Applied Poultry
Research, -: +0. +33..
Harper AE, Benevenga NJ and Wohlhueter RM. E#ects of
ingestion of disproportionate amounts of amino acids. Physiological Reviews, /*: .,2//2. +31*.
Hasegawa H, Shinohara Y, Akahane K and Hashimoto T. Direct
detection and evaluation of conversion of D-methionine into
L-methionine in rats by stable isotope methodology. Journal
of Nutrition, +-/: ,**+,**/. ,**/.
Hoehler D and Hooge DM. Relative e#ectiveness of methionine
sources in turkeys: scientic and new commercial data. International Journal of Poultry Science, ,: -0+-00. ,**-.
Huyghebaert G. Comparison of DL-methionine and methionine
hydroxy analogue: free acid in broilers by using multiexponential regression model. British Poultry Science, -.:

177

-/+-/3. +33-.
Huyghebaert G and Pack M. E#ects of dietary protein content,
addition of nonessential amino acids and dietary methionine
to cysteine balance on responses to dietary sulphurcontaining amino acids in broilers. British Poultry Science,
-1: 0,-0-3. +330.
Kim WK, Froelich CA Jr, Patterson PH and Ricke SC. The
potential to reduce poultry nitrogen emissions with dietary
methionine or methionine analogues supplementation.
Worlds Poultry Science Journal, 0,: --2-/-. ,**0.
Knight CD and Dibner JJ. Comparative absorption of ,hydroxy-.-(methylthio)-butanoic acid and L-methionine in
the broiler chick. Journal of Nutrition, ++.: ,+13,+20.
+32..
Knight CD, Wuelling CW, Atwell CA and Dibner JJ. E#ect of
intermittent periods of high environmental temperature on
broiler performance responses to sources of methionine activity. Poultry Science, 1-: 0,10-3. +33..
Leclercq B, Chaqneau AM, Cochard T, Hamaoui S and Larbire
M. Comparative utilisation of sulphur-containing amino
acids by genetically lean or fat chickens. British Poultry
Science, -.: -2--3+. +33-.
Lemme A, Hoehler D, Brennan JJ and Mannion PF. Relative
e#ectiveness of methionine hydroxy analog compared to
DL-methionine in broiler chickens. Poultry Science, 2+:
2-22./. ,**,.
Lerner J, Sattelmeyer P and Rush R. Kinetics of methionine
inux into various regions of chicken intestine. Comparative
Biochemistry and Physiology, /*A: ++-+,*. +31/.
Lingens G and Molnar S. Studies on metabolism of broilers by
using +.C-labelled DL-methionine and DL-methionine
hydroxy analogue Ca-salt. Archiv fur Tierernahrung, .3:
++-+,.. +330.
Liu Z, Bateman A, Bryant M, Abebe A and Roland D. Estimation of bioavailability of DL-methionine hydroxy analogue
relative to DL-methionine in layers with exponential and
slope-ratio model. Poultry Science, 2-: +/2*+/20. ,**..
Maenz DD and Engele-Schaan CM. Methionine and ,-hydroxy.-methylthiobutanoic acid are transported by distinct Na
-dependent and H-dependent in the brush border membrane of the chick intestinal epithelium. Journal of Nutrition, +,0: /,3/-0. +330.
Mart
n-Venegas R, Geraert PA and Ferrer R. Conversion of the
methionine hydroxy analogue DL-,-hydroxy-(.-methylthio)
butanoic acid to sulfur containing amino acids in the chicken small intestine. Poultry Science, 2/: +3-,+3-2. ,**0.
Matsushita K, Takahashi K and Akiba Y. E#ects of adequate or
marginal excess of dietary methionine hydroxy analogue
free acid on growth performance, edible meat yields and
inammatory response in female broiler chickens. Journal
of Poultry Science, ..: ,0/,1,. ,**1.
McDevitt RM, Mack S and Wallis IR. Can betaine partially
replace or enhance the e#ect of methionine by improving
broiler growth and carcase characteristics? British Poultry
Science, .+: .1-.2*. ,***.
Mendonca CX and Jensen LS. Inuence of protein concentration
on the sulphur-containing amino acid requirement of broiler
chickens. British Poultry Science, -*: 223232. +323.
Mitchell MA and Lemme A. Examination of the composition of
the luminal uid in the small intestine of broilers and absorption of amino acids under various ambient temperatures
measured in vivo. International Journal of Poultry Science,

178

J. Poult. Sci., .0 (-)

1: ,,-,--. ,**2.
Molitoris BA and Baker DH. The choline requirement of broiler
chicks during the seventh week of life. Poultry Science, //:
,,*,,.. +310.
Morris TR and Gous RM. Partitioning of the response to protein
between egg number and egg weight. British Poultry Science, ,3: 3-33. +322.
Morris TR, Gous RM and Abebe S. E#ects of dietary protein
concentration on the response of growing chicks to
methionine. British Poultry Science, --: 13/2*-. +33,.
Noll SL, Waibel PE, Cook DR and Witmer JA. Biopotency of
methionine sources for young turkeys. Poultry Science, 0-:
,./2,.1*. +32..
Novak C, Yakout H and Scheideler S. The combined e#ects of
dietary lysine and total sulfur amino acid level on egg
production parameters and egg components in Dekalb Delta
laying hens. Poultry Science, 2-: 31132.. ,**..
Noy Y and Sklan D. Uptake capacity in vitro for glucose and
methionine and in situ for oleic acid in the proximal small
intestine of posthatch chicks. Poultry Science, 1/: 332+**,.
+330.
NRC. Nutrient Requirement for Poultry, 3th Ed. National
Academy Press, Washington DC, USA. +33..
Pan Y, Wong EA, Dibner JJ, Va
zquez-An

on M and Webb Jr
KE. Poly(A) RNA encoding proteins capable of transporting L-methionine and/or DL-,-hydroxy-.-(methylthio)
butanoic acid are present in the intestinal mucosa of broilers.
Journal of Nutrition, +-,: -2,-20. ,**,.
Peterson CF, Sauter EA, Steele EE and Parkinson JF. Use of
methionine intake restriction to improve egg shell quality by
control of egg weight. Poultry Science, 0,: ,*..,*.1. +32-.
Picard ML, Uzu G, Dunnington EA and Siegel PB. Food intake
adjustments of chicks: short term reactions to deciencies in
lysine, methionine and tryptophan. British Poultry Science,
-.: 1-11.0. +33-.
Pillai PB, Fanatico AC, Beers KW, Blair ME and Emmert JL.
Homocysteine remethylation in young broilers fed varying
levels of methionine, choline, and betaine. Poultry Science,
2/: 3*3/. ,**0.
Plavnik Y and Bornstein S. The sparing action of inorganic
sulphate on sulphur amino acids in practical broiler diets:
The replacement of some of the supplementary methionine
in broiler nisher diets. British Poultry Science, +3: +/3
+01. +312.
Poeikhampa T. E#ect of dietary protein and methionine level on
laying performance and chemical body composition of
laying hen raised in closed house system. Master Thesis,
Kasetsart University, pp. 3*. ,**..
Poosuwan K, Bunchasak C and Kaewtapee C. Long-term feeding e#ects of dietary protein levels on egg production, immunocompetence and plasma amino acids of laying hens in
subtropical condition. Journal of Animal Physiology and
Animal Nutrition. ,**3 (in press).
Poultry Research Foundation, Annual Report, The University of
Sydney, Sydney, ,***.
Pourreza J and Smith WK. Performance of laying hens fed on
low sulphur amino acids diets supplemented with choline
and methionine. British Poultry Science, ,3: 0*/0++. +322.
Ravikiran D and Devegowda G. E#ects of DL-methionine
supplementation in the ration of commercial layers during
summer. Indian Journal of Poultry Science, --: ,13,2-.
+332.

Reid BL, Madrid A and Maiodrino RM. Relative biopotency of

three methionine sources for laying hens. Poultry Science,
0+: 1,01-*. +32,.
Ribeiro AML, Dahlke F and Kessler AM. Methionine sources do
not a#ect performance and carcass yield of broilers fed
vegetable diets and submitted to cyclic heat stress. Brazilian
Journal of Poultry Science, 1: +/3+0.. ,**/.
Rostagno HS and Barbosa WA. Biological e$cacy and absorption of DL-methionine hydroxy analogue free acid compared to DL-methionine in chickens as a#ected by heat
stress. British Poultry Science, -0: -*--+,. +33/.
Rostagno HS, Pupa JMR and Pack M. Diet formulation for
broilers based on total versus digestible amino acid. Journal
of Applied Poultry Research, .: ,3-,33. +33/.
Rostagno HS and Pack M. Can betaine replace supplemental
DL-methionine in broiler diets? Journal of Applied Poultry
Research, /: +/*+/.. +330.
Sasse CE and Baker DH. Sulfur utilization by the chick with
emphasis on the e#ect of inorganic sulfate on the cystinemethionine interrelationship. Journal of Nutrition, +*.:
,..,/+. ,***.
Sauer N, Emrich K, Piepho HP, Lemme A, Redshaw MS and
Mosenthin R. Meta-analysis of the relative e$ciency of
methionine-hydroxy-analogue-free-acid compared with DLmethionine in broilers using nonlinear mixed models. Poultry Science, 21: ,*,-,*-+. ,**2.
Schutte JB and van Weerden EJ. Requirement of the hen for
sulphur-containing amino acids. British Poultry Science, +3:
/1-/2+. +312.
Schutte JB, van Weerden EJ and Bertram HL. Sulphur amino
acid requirement of laying hens and the e#ects of excess
dietary methionine on laying performance. British Poultry
Science, ,.: -+3-,0. +32-.
Schutte JB, De Jong J and Bertram HL. Requirement of the
laying hen for sulfur amino acids. Poultry Science, 1-: ,1.
,2*. +33..
Schutte JB and Pack M. Sulfur amino acid requirement of broiler
chicks from fourteen to thirty-eight days of age. +. Performance and carcass yield. Poultry Science, 1.: .2*.21. +33/.
Schutte JB, De Jong J, Smink W and Pack M. Replacement value
of betaine for DL-methionine in male broiler chicks. Poultry
Science, 10: -,+-,/. +331.
Scott ML. Studies on the comparative utilization of synthetic
sources of methionine activity in laying pullets. Nutrition
Reports International, -0: +*.-+*/,. +321.
Sekiz SS, Scott ML and Nesheim MC. The e#ect of methionine
deciency on body weight, food and energy utilization in the
chick. Poultry Science, /.: ++2.++22. +31/.
Sell DR and Rogler JC. The e#ect of sorghum tannin and
methionine on the performance of laying hens maintained in
two temperature environments. Poultry Science, 0-: +*3
++0. +32-.
Shafer DJ, Carey JB, Prochaska JF and Sams AR. Dietary
methionine intake e#ects on egg component yield, composition, functionality, and texture prole analysis. Poultry Science, 11: +*/0+*0,. +332.
Silva RGC Jr, Lana GRQ, Rabello CBV and Boa-Viagem C.
Requirements of methionine cystine for female broilers
chickens from + to ,+ and ,, to ., days old on tropical
climate region. Revista Brasileira de Zootecnia, -/: .31
/*-. ,**0.
Soriano-Garc
a JF, Torras-Llort M, Ferrer R and Moret
o M.

Bunchasak: Methionine for Poultry Production

Multiple pathways for L-methionine transport in brushborder membrane vesicles from chicken jejunum. The Journal of Physiology, /*3: /,1/-3. +332.
Soriano-Garc
a JF, Torras-Llort M, Moreto
M and Ferrer R.
Regulation of L-methionine and L-lysine uptake in chicken
jejunal brush-border membrane by dietary methionine. The
American Journal of Physiology - Regulatory, Integrative
and Comparative Physiology, ,11: R+0/.R+00+. +333.
Sterling KG, Pesti GM and Bakalli RI. Performance of di#erent
broiler genotypes fed diets with varying levels of dietary
crude protein and lysine. Poultry Science, 2/: +*./+*/..
,**0.
Summers JD, Atkinson JL and Spratt D. Supplementation of a
low protein diet in an attempt to optimize egg mass output.
Canadian Journal of Animal Science, 1+: ,++,,*. +33+.
Summers JD, Spratt D and Atkinson JL. Broiler weight gain and
carcass composition when fed diets varying in amino acid
balance, dietary energy, and protein level. Poultry Science,
1+: ,0-1-. +33,.
Swain BK and Johri TS. E#ect of supplemental methionine,
choline and their combinations on the performance and
immune response of broilers. British Poultry Science, .+:
2-22. ,***.
Takahashia K, Ohta N and Akiba Y. Inuences of dietary
methionine and cysteine on metabolic responses to immunological stress by Escherichia coli lipopolysaccharide injection, and mitogenic response in broiler chickens. British
Journal of Nutrition, 12: 2+/2,+. +331.
Torras-Llort M, Ferrer R, Soriano-Garc
a JF and Moreto
M.
L-Lysine transport in chicken jejunal brush border membrane vesicles. Journal of Membrane Biology, +/,: +2-+3-.
+330.
Torras-Llort M, Soriano-Garc
a JF, Ferrer R and Moreto
M.
E#ect of a lysine-enriched diet on L-lysine transport by the
brush border membrane of the chicken jejunum. American
Journal of Physiology, ,1.: R03R1/. +332.
Tsiagbe VK, Cook ME, Harper AE and Sunde ML. Enhanced
immune responses in broiler chicks fed methioninesupplemented diets. Poultry Science, 00: ++.1++/.. +321.
van Heugten E, Spears JW, Co#ey MT, Kegley EB and Qureshi
MA. The e#ect of methionine and aatoxin on immune
function in weanling pigs. Journal of Animal Science, 1,:
0/200.. +33..
van Weerden EJ and Schutte JB. Comparison of DL-methionine,
DL-methionine-Na, DL-methionine hydroxy analogue-Ca,
and DL-methionine hydroxy analogue-free acid with layers.
Poultry Science, 0-: +13-+133. +32..
Va
zquez-An

on M, Gonza
lez-Esquerra R, Saleh E, Hampton T,

179

Ritcher S, Firman J and Knight CD. Evidence for

,-hydroxy-.(methylthio) butanoic acid and DL-methionine
having di#erent dose responses in growing broilers. Poultry
Science, 2/: +.*3+.,*. ,**0.
Veltmann JR Jr, Wyatt RD and Voight MN. The e#ect of
varying the total sulfur amino acid content in the diets of
chicks with aatoxicosis. Poultry Science, 0*: +1.2 (Abstract). +32+.
Veltmann JR Jr, Wyatt RD, Voight MN and Shamsuddin Z.
Inuence of dietary sulfur amino acid levels on performance, free amino acids and biochemical parameters in
plasma and hepatic glutathione of broiler chicks fed
aatoxin. Poultry Science, 0,: +/+2 (Abstract). +32-.
Vieira SL, Lemme A, Goldenberg DB and Brugalli I. Responses
of growing broilers to diets with increased sulfur amino
acids to lysine ratios at two dietary protein levels. Poultry
Science, 2-: +-*1+-+-. ,**..
Waldroup PW, Mitchell RI, Payne JR and Hazen KR. Performance of chicks fed diets formulated to minimize excess levels
of essential amino acids. Journal of Poultry Science, //:
,.-,/-. +310.
Waldroup PW. Inuence of environmental temperature on protein and amino acid needs of poultry. Federation Proceeding, .+: ,2,+,2,-. +32,.
Waldroup PW and Hellwig HM. Methionine and total sulfur
amino acid requirements inuenced by stage of production.
Journal of Applied Poultry Research, .: ,2-,3,. +33/.
Wallis IR. Dietary supplements of methionine increase breast
meat yield and decrease abdominal fat in growing broiler
chickens. Australian Journal of Experimental Agriculture,
-3: +-++.+. +333.
Xie MS, Hou S, Huang W and Fan HP. E#ect of excess
methionine and methionine hydroxy analogue on growth
performance and plasma homocysteine of growing Pekin
ducks. Poultry Science, 20: +33/+333. ,**1.
Yalcin S, Ozkan S, Acikgoz Z and Ozkan K. E#ect of dietary
methionine on performance, carcase characteristics and
breast meat composition of heterozygous naked neck (Na/
na) birds under spring and summer conditions. British
Poultry Science, .*: 02203.. +333.
Zhan XA, Li JX, Xu ZR and Zhao RQ. E#ects of methionine
and betaine supplementation on growth performance, carcass composition and metabolism of lipids in male broilers.
British Poultry Science, .1: /10/2*. ,**0.
Zhang LB and Guo YM. E#ects of liquid DL-,-hydroxy-.-methylthio
butanoic acid on growth performance and immune responses in broiler chickens. Poultry Science, 21: +-1*+-10.
,**2.