2006 Poultry Science Association, Inc.

Effects of Betaine and Choline on Response

to Methionine Supplementation to Broiler Diets
Formulated to Industry Standards1

P. W. Waldroup,2 M. A. Motl, F. Yan, and C. A. Fritts

Department of Poultry Science, University of Arkansas, Fayetteville 72701

Primary Audience: Nutritionists, Veterinarians, Poultry Processors, Researchers

SUMMARY
A study was conducted to evaluate the Met-sparing effects of the methyl donors, choline (CHO)
and betaine (BET). Male broilers of a commercial strain were fed corn-soybean meal-based diets
formulated to meet nutritional levels typical of top broiler producers. Diets were fortified with a
complete vitamin premix devoid of CHO. In test diets, Met levels were as formulated, or less 0.05,
0.10, 0.15, or 0.20% Met. Within each Met level, diets received no supplemental CHO or BET or
1,000 mg of CHO/kg, 1,000 mg of BET/kg, or a combination of 500 mg each of CHO and BET/
kg. This resulted in a 4 × 5 factorial arrangement with each of the 20 treatments fed to 4 pens of
60 birds for a 56-d feeding trial. Samples of birds were processed at 42, 49, and 56 d to determine
parts yield and dressing percentage. Intestinal segments were evaluated for tensile strength at
these same ages.
The dietary Met level had no significant impact on BW at 14 or 56 d of age, but at 35, 42, and
49 d reduction of the Met content resulted in loss of BW. Feed conversion was not affected by the
Met level at 14 d but was significantly affected by reducing the Met level at all other ages. Breast
meat yield was reduced in a linear manner by reductions in Met supplementation. The CHO or
BET supplementation had no apparent sparing effect on Met needs but did improve feed conversion
at 35 and 42 d. There was a positive effect of CHO and BET on breast yield that was independent
of Met levels; CHO was as effective as BET for this purpose. No effect of CHO or BET on intestinal
strength was observed in the present study.

Key words: broiler, choline, betaine, methionine, sparing effect

2006 J. Appl. Poult. Res. 15:58–71

DESCRIPTION OF PROBLEM then to homocysteine. The back reaction requires

Methionine is the first limiting amino acid
in broiler diets based on soybean meal, and it
functions as a key intermediate in methyl group
transfer. Numerous methylation reactions in-
volve Met, in which Met is converted to S-aden-
osylmethionine, the methyl group donor, and
betaine (BET) or tetrahydrofolate as the pro-
vider of the methyl group necessary to convert
homocysteine to Met [1, 2]. Both Met and homo-
cysteine are potentially toxic to cells, and both
may accumulate if methyl donors or methyl ac-
ceptors are deficient. Alternative methyl donors
may alleviate the dietary needs for Met by taking

1
Published with the approval of the Director, Arkansas Agricultural Experiment Station, Fayetteville, AR 72701. Mention
of a trade name, proprietary product, or specific equipment does not constitute a guarantee or warranty by the University
of Arkansas and does not imply its approval to the exclusion of other products that may be suitable.
2
Corresponding author: Waldroup@uark.edu
WALDROUP ET AL.: INDUSTRY BROILER DIETS 59

Table 1. Nutrient analysis of diets from the top 5 broiler companies in an agricultural survey

Starter Grower Finisher 1 Finisher 2

Nutrient 0–14 d 14–35 d 35–42 d 42–56 d

ME, kcal/lb 1,362.00 1,394.00 1,416.00 1,460.00

Crude protein, % 20.93 19.16 17.02 16.08
Lys, % 1.23 1.09 0.93 0.88
TSAA, % 0.96 0.85 0.76 0.76
Trp, % 0.25 0.22 0.20 0.18
Arg, % 1.52 1.31 1.17 1.08
Thr, % 0.85 0.76 0.68 0.64
Nonphytate P, % 0.43 0.37 0.34 0.33
Calcium, % 0.83 0.85 0.76 0.76
Sodium, % 0.23 0.20 0.21 0.20

the place of Met as a methyl donor per se or by
providing the methyl group necessary for the
conversion of homocysteine to Met.
Choline (CHO) has long been used as a
vitamin supplement for broiler diets. Numerous
studies have investigated the interrelationship
between CHO and Met as donors of methyl
groups, with considerable variation in results [3,
4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14]. More recently,
BET has been evaluated as a methyl donor as a
means of reducing the needs for Met [15, 16,
17, 18, 19, 20], also with variation in results.
The present trial was conducted to examine the
potential value of CHO and BET, alone or in
combination, in reducing the dietary needs for
Met in broiler diets.
MATERIALS AND METHODS
Nutritional specifications for the top 5 com-
panies participating in a popular agricultural sur-
vey [21] were used as the basis of formulation
(Table 1). Diets were formulated using corn and
soybean meal of known composition and supple-
mental amino acids to meet exactly the specified
requirements for CP, ME, Ca, available P, Na,
Lys, TSAA, Arg, and Thr with only slight ex-
cesses of Try (Table 2). Feeding intervals were
0 to 14 d for starter, 14 to 35 d for grower, 35
to 42 d for finisher 1, and 42 to 56 d for finisher
2, similar to the intervals used by the top 5 com-
panies.
The amount of added Met in the diets as
formulated was 0.30% for starter, 0.24% for
grower, 0.20% for finisher 1, and 0.22% for
finisher 2. For test diets, Met levels were 1) as
formulated, 2) less 0.05%, 3) less 0.10%, 4)
less 0.15%, and 5) less 0.20%. Within each Met
level, diets received 1) no supplemental CHO
or BET, 2) 1,000 mg of CHO/kg, 3) 1,000 mg
of BET/kg, or 4) 500 mg of CHO + 500 mg of
BET/kg. The CHO was supplied as a dry 60%
choline chloride product [22], whereas BET was
added as a crystalline 97% product [23]. This
resulted in a 4 × 5 factorial arrangement of treat-
ments. A common basal diet was mixed within
each test period, and aliquots were used to mix
each of the test diets. All diets contained monen-
sin at 90 g/ton as an anticoccidial. Each treat-
ment was fed to 4 pens of 60 male chicks. The
basal diet within each feeding period was evalu-
ated for total amino acids, and all test diets were
analyzed for supplemental MET, CHO, and BET
by commercial laboratories.
Day-old male chicks of a commercial strain
[24] were obtained from a local hatchery where
they had been vaccinated in ovo for Marek’s
disease and had received vaccinations for new-
castle disease and infectious bronchitis post-
hatch via a coarse spray. They were randomly
distributed among 80 pens in a curtain-sided
broiler house of commercial design. Sixty birds
were placed in each pen. New softwood shavings
over concrete floors served as litter. Each pen
was equipped with 2 tube feeders and 1 auto-
matic water fount. Supplemental feeders and wa-
terers were used during the first 7 d. Fluorescent
lights supplemented natural daylight to provide
23 h light daily. Automatic brooder stoves and
fans controlled temperature and airflow. Care
and management of the birds followed recom-
mended guidelines [25].
Group BW by pen was obtained at 14, 35,
42, 49, and 56 d. Feed consumption was deter-
mined at each feed change interval. Birds were
60 JAPR: Research Report

Table 2. Composition (g/kg) of diets formulated to nutrient standards of top broiler companies

Starter Grower Finisher 1 Finisher 2

Ingredient 0–14 d 14–35 d 35–42 d 42–56 d

Yellow corn 621.21 655.37 718.36 722.44

Soybean meal (48% CP) 312.75 276.01 218.11 198.36
Dicalcium phosphate 20.39 17.27 15.27 14.77
Poultry oil 16.67 23.32 20.69 36.73
Ground limestone 8.70 10.86 10.16 10.42
Feed-grade salt 5.24 4.67 4.76 4.68
BMD-501 0.50 0.50 0.50 0.50
Coban-602 0.75 0.75 0.75 0.75
Vitamin premix3 2.00 2.00 2.00 2.00
Trace mineral mix4 1.00 1.00 1.00 1.00
L-Lys HCl (98%) 1.88 1.27 1.30 1.29
L-Thr 0.90 0.57 0.65 0.64
L-Arg 1.93 1.02 1.39 1.16
DL-Met (98%) 3.08 2.39 2.06 2.26
Variable5 3.00 3.00 3.00 3.00
Total 1,000.00 1,000.00 1,000.00 1,000.00
Calculated analysis
ME, kcal/lb 1,362.00 1,394.00 1,416.00 1,460.00
Crude protein, % 20.93 19.17 17.02 16.08
Lys, % 1.24 1.09 0.93 0.88
TSAA, % 0.96 0.85 0.76 0.76
Met, % 0.63 0.53 0.48 0.49
Trp, % 0.27 0.24 0.20 0.19
Arg, % 1.52 1.31 1.17 1.08
Thr, % 0.85 0.76 0.68 0.64
Nonphytate P, % 0.43 0.37 0.34 0.33
Calcium, % 0.84 0.85 0.76 0.75
Sodium, % 0.23 0.20 0.21 0.20
1
Alpharma, Inc., Fort Lee, NJ.
2
Elanco Animal Health division of Eli Lilly and Co., Indianapolis, IN.
3
Provided per kilogram of diet: vitamin A (from vitamin A acetate), 7,714 IU; cholecalciferol, 2,204 IU; vitamin E (from
DL-α-tocopheryl acetate), 16.53 IU; vitamin B12, 0.013 mg; riboflavin, 6.6 mg; niacin, 39 mg; pantothenic acid, 10 mg;
menadione (from menadione dimethyl-pyrimidinol), 1.5 mg; folic acid, 0.9 mg; thiamin (from thiamin mononitrate), 1.54
mg; pyridoxine (from pyridoxineⴢHCl), 2.76 mg; D-biotin, 0.066 mg; ethoxyquin, 125 mg; Se, 0.1 mg.
4
Provided per kilogram of diet: Mn (from MnSO4ⴢH2O), 100 mg; Zn (from ZnSO4ⴢ7H2O), 100 mg; Fe (from FeSO4ⴢ7H2O),
50 mg; Cu (from CuSO4ⴢ5H2O), 10 mg; I from Ca(IO3)2ⴢH2O), 1 mg.
4
Variable amounts of choline or betaine.

checked twice daily and weight of dead birds
used to adjust feed conversion. At 42, 49, and
56 d, 5 birds per pen were randomly selected
for processing from among 20 birds marked at
1 d of age by a toe slit. After a 12-h fast, the
birds were placed in coops and transported ap-
proximately 1 mi to the processing plant. They
were weighed and processed with automated
evisceration as described by Fritts and Waldroup
[26]. Due to the automatic evisceration it was
not possible to determine abdominal fat content.
The eviscerated carcass was weighed and placed
in a chiller with air agitation for 1 h. The car-
casses were removed, hung on shackles, and
allowed to drain for 30 min. The carcasses were
then deboned by trained technicians to determine
parts yield (breast, leg quarters, and wings).
Parts yield was expressed as percentage of evis-
cerated chilled carcass.
At 42, 49, and 56 d, intestinal segments were
obtained from the slaughtered birds and intesti-
nal breaking strength and elasticity determined
as described by Huff et al. [27]. The test instru-
ment used for determining the breaking force of
the intestines was a Chatillon model DFM-10
digital break force meter [28] using a 4-in. seg-
ment of intestine with a speed set at 10 in./min.
Pen means served as the experimental unit.
Data were subjected ANOVA as a 4 × 5 factorial
arrangement of treatments using the GLM option
WALDROUP ET AL.: INDUSTRY BROILER DIETS 61

Table 3. Total and added choline content of test diets (mg/kg)

Added
choline Starter Grower Finisher 1 Finisher 2
(mg/kg) Basis1 0–14 d 14–35 d 35–42 d 42–56 d

0 C 1,239 1,160 1,051 981

A 1,193 1,116 1,024 925
+500 D 417 414 420 418
+1,000 D 748 768 762 756
1
C = calculated from NRC (1994); A = analyzed by Danisco Animal Nutrition, St. Louis, MO; D = difference between
analyzed value and amount provided by the unsupplemented diet.

of SAS [29]. The main effects of level of supple- BET averaged 544 and 1,075 mg/kg, respec-
mental Met and the various additives or combi- tively. Low levels of BET were found in the
nations served as main effects; the interaction basal diets; levels from 35 to 42 d were below
of Met levels and the various additives was also assay detection limits, perhaps due to different
evaluated. When statistical significance among sources of corn and soybean meal. Calculated
or between treatment means was observed, data and analyzed levels of supplemental Met are
were separated using the least square means op- shown in Table 5. Analyzed levels were in
tion of SAS [29]. Selected data were subjected close agreement with calculated levels over the
to regression analysis using the PROCREG op- entire range of test diets. Thus, the test diets
tion of SAS. Statements of probability are based were considered to contain the anticipated lev-
on P < 0.05 unless stated otherwise. els of CHO, BET, and Met.
The effect of levels of CHO, BET, and Met
RESULTS on BW at different ages is shown in Table 6.
Total and added CHO contents of mixed When compared by ANOVA, the Met content
feeds are listed in Table 3. Calculated values of the diet had no significant effect on BW at
for CHO in the basal diet were based on NRC 14 or 56 d of age. At 35 and 42 d of age,
[30]. Added CHO levels were determined by however, reduction of Met to 0.15 and 0.10%,
subtracting total CHO content of the diets with respectively, less than that in the commercial
no added CHO from that of diets with 500 or diets significantly reduced BW compared with
1,000 mg of CHO/kg added. Analyzed levels the control group. At 49 d, a reduction of more
for the unsupplemented basal diet were slightly than 0.05% resulted in a significant reduction
less than calculated; added levels for the 500 in BW. However, when subjected to regression
and 1,000 mg/kg levels over all age periods analysis, a significant linear regression of BW
averaged approximately 417 and 758 mg/kg, on level of Met was observed at all ages. For
respectively. Analyzed BET content of the each 0.01% reduction in level of Met, there
mixed feeds is shown in Table 4. Total BET were BW reductions of 0.68, 5.44, 7.08, 5.74,
levels in diets with 500 and 1,000 mg/kg added and 4.72 g at 14, 35, 42, 49, and 56 d of age,
respectively (R2 of 0.79, 0.85, 0.86, 0.97, and
0.91, respectively). Numerical improvements
Table 4. Analyzed betaine content of test diets (mg/
kg)1 in BW were frequently observed when CHO
or BET was added alone or in combination;
Added betaine (mg/kg) however, these did not reach a level of statisti-
Feeding Age
period fed None 500 1,000 cal significance (P < 0.05). There was no inter-
action between Met level and the addition of
Starter 0–14 d 81 560 1,132
Grower 14–35 d 80 546 1,121 CHO or BET, alone or in combination.
Finisher 1 35–42 d BDL2 444 916 The effect of levels of CHO, BET, and Met
Finisher 2 42–56 d 77 625 1,130 on feed conversion at different ages is shown
1
Analyzed by Danisco Animal Nutrition, St. Louis, MO. in Table 7. No significant effect of Met on
2
Below detection limits. feed conversion was observed at 14 d; however,
62 JAPR: Research Report

Table 5. Calculated and analyzed level of supplemental Met in test diets (% of diet)

Reduction
from Starter Grower Finisher 1 Finisher 2
standard (%) Basis1 0–14 d 14–35 d 35–42 d 42–56 d

0 C 0.310 0.241 0.208 0.228

A 0.304 0.228 0.192 0.218
−0.05 C 0.259 0.191 0.158 0.178
A 0.267 0.170 0.145 0.182
−0.10 C 0.209 0.140 0.107 0.127
A 0.215 0.139 0.098 0.123
−0.15 C 0.158 0.089 0.056 0.077
A 0.152 0.079 0.051 0.065
−0.20 C 0.108 0.039 0.008 0.026
A 0.111 0.037 0.013 0.025

C = calculated level; A = analyzed by Degussa Corporation, Allendale, NJ.

1

cumulative feed conversion at 35, 42, 49, and
56 d was significantly influenced by Met level
of the diet. In general, a reduction of up to
0.10% below the industry average Met level
could be achieved without a significant change
in feed conversion. Greater reductions in Met
had a detrimental effect. Regression analysis
indicated a quadratic response to levels of Met,
with a breakpoint at approximately 0.10% re-
moval of Met at 35, 42, 49, and 56 d (Figure
1). The addition of 1,000 mg of CHO/kg, 1,000
mg of BET/kg, or the combination of 500 mg
of each significantly improved feed conversion
over the birds fed the unsupplemented diets at
35, 42, 49, and 56 d of age with little difference
among these 3 treatments. At 14 d, there was
a significant interaction of CHO and BET sup-
plementation with level of Met. However, this
followed no consistent pattern and was not ob-
served at other ages, although it neared signifi-
cance at 42 (P = 0.08) and 49 (P = 0.06) d
of age.
No significant differences among or be-
tween dietary treatments were observed for
mortality (Table 8). Birds were grown on new
softwood shavings and received an anticoccid-
ial and a growth-promoting antibiotic through-
out the duration of the study. Mortality was
within normal limits for our facility.
Dietary treatments had no significant ef-
fects on dressing percentage of birds at 42, 49,
or 56 d of age (Table 9). Because fat deposits
contained in the viscera were removed from
the carcass by the automatic evisceration, it
was not possible to determine the amount of
abdominal fat. There were no indications of
interaction of Met levels and the addition of
CHO or BET, alone or in combination.
Significant dietary effects on breast yield
at 42, 49, and 56 d of age were observed (Table
10). Reduction of Met levels below that of the
top industry producers resulted in significant
reduction in breast yield that followed a linear
trend. Each reduction of 0.01% in Met level
from that used by the top industry producers
resulted in losses of 0.08, 0.12, and 0.08% of
breast meat yield at 42, 49, and 56 d, respec-
tively (R2 of 0.92, 0.92, and 0.83, respectively).
Addition of 1,000 mg of CHO/kg, 1,000 mg of
BET/kg, or a combination of 500 mg of each
to the diets resulted in significant improve-
ments in breast yield at 42, 49, and 56 d of
age. This response was independent of level of
Met in the diet, as evidenced by lack of an
interaction of Met and the 2 supplements. No
differences between the response to CHO or
BET and the combination of the 2 was observed
at 42 or 49 d; at 56 d the breast yield of birds
fed BET alone was superior to that of birds
fed CHO alone, with the combination of the 2
intermediate between the 2 supplements.
Tensile strength of intestinal segments is
shown in Table 11. There was no significant
effect of levels of Met or of the addition of CHO
or BET, alone or in combination. As mentioned
previously an anticoccidial was fed for the du-
ration of the study, and a growth promoting
antibiotic known to be effective against clos-
tridial infection was also included in all diets.
Periodic examination of birds that died during
WALDROUP ET AL.: INDUSTRY BROILER DIETS 63

Table 6. Effects of levels of methionine, choline, and betaine on body weight of male broilers at different ages
(means of 4 pens of 60 birds/treatment)

500 mg
of choline/kg
Reduction No choline 1,000 mg 1,000 mg 500 mg
in Met (%) or betaine of choline/kg of betaine/kg of betaine/kg Average

14 d BW (g)
0.00 346 336 340 323 336
0.05 334 334 344 329 335
0.10 338 307 342 320 326
0.15 316 322 320 333 323
0.20 332 329 330 312 325
Average 333 325 335 323
35 d BW (g)
0.00 1,882 1,859 1,887 1,799 1,857ab
0.05 1,846 1,891 1,935 1,841 1,878a
0.10 1,847 1,790 1,819 1,839 1,824ab
0.15 1,748 1,785 1,824 1,851 1,802bc
0.20 1,731 1,757 1,802 1,747 1,759c
Average 1,811 1,816 1,854 1,815
42 d BW (g)
0.00 2,474 2,493 2,460 2,413 2,460ab
0.05 2,480 2,496 2,497 2,438 2,478a
0.10 2,395 2,413 2,401 2,421 2,407bc
0.15 2,337 2,385 2,407 2,448 2,394c
0.20 2,246 2,345 2,376 2,333 2,325d
Average 2,386 2,426 2,428 2,411
49 d BW (g)
0.00 2,981 2,992 2,955 2,964 2,973a
0.05 2,906 2,956 3,003 2,930 2,949ab
0.10 2,892 2,875 2,931 2,906 2,901bc
0.15 2,816 2,889 2,894 2,943 2,886bc
0.20 2,805 2,892 2,886 2,863 2,861c
Average 2,880 2,921 2,934 2,921
56 d BW (g)
0.00 3,380 3,389 3,381 3,427 3,394
0.05 3,308 3,381 3,434 3,353 3,369
0.10 3,337 3,299 3,358 3,309 3,326
0.15 3,296 3,328 3,240 3,364 3,307
0.20 3,239 3,345 3,390 3,252 3,307
Average 3,312 3,348 3,361 3,341
0–14 d 0–35 d 0–42 d 0–49 d 0–56 d
Source of
variance P diff SEM P diff SEM P diff SEM P diff SEM P diff SEM

Met level 0.45 6 0.001 20 0.0001 20 0.01 24 0.24 33

Supplements 0.34 6 0.37 19 0.37 18 0.40 23 0.72 30
Met × supplements 0.81 13 0.62 40 0.52 40 0.94 50 0.82 65
CV 7.94 4.83 3.50 3.63 4.13
Means within an age with common superscripts do not differ significantly (P < 0.05).
a–d

the study by a qualified poultry veterinarian
revealed no indication of coccidiosis.
DISCUSSION
Results of the present study indicate that
reduction in Met and TSAA levels from those
currently used by the top broiler producers may
result in loss of live performance and in breast
meat yield. Reduction of more than 0.10% from
current values could be tolerated without ad-
verse effects on BW gain or feed conversion;
however, breast meat yield was negatively af-
64 JAPR: Research Report

Table 7. Effects of levels of Met, choline, and betaine on feed conversion by male broilers at different ages (means
of 4 pens of 60 birds/treatment)

500 mg
of choline/kg
Reduction No choline 1,000 mg 1,000 mg 500 mg
in Met (%) or betaine of choline/kg of betaine/kg of betaine/kg Average

0–14 d feed:gain ratio

0.00 1.133bcd 1.154abc 1.173ab 1.163abc 1.156
0.05 1.186a 1.138bc 1.142abc 1.131cd 1.149
0.10 1.132bcd 1.136bcd 1.149abc 1.139bc 1.139
0.15 1.166abc 1.148abc 1.136bcd 1.099d 1.137
0.20 1.141bc 1.167abc 1.148abc 1.127cd 1.146
Average 1.151 1.149 1.150 1.132
0–35 d feed:gain ratio
0.00 1.523 1.520 1.562 1.513 1.530bc
0.05 1.532 1.496 1.515 1.500 1.511c
0.10 1.539 1.508 1.527 1.499 1.518d
0.15 1.581 1.528 1.523 1.531 1.541b
0.20 1.627 1.599 1.564 1.570 1.590a
Average 1.561x 1.530y 1.538y 1.523y
0–42 d feed:gain ratio
0.00 1.666 1.649 1.720 1.654 1.672bc
0.05 1.673 1.648 1.664 1.648 1.658c
0.10 1.700 1.646 1.667 1.651 1.666c
0.15 1.716 1.672 1.677 1.682 1.687b
0.20 1.793 1.747 1.724 1.720 1.746a
Average 1.710x 1.673yz 1.690y 1.671z
0–49 d feed:gain ratio
0.00 1.767 1.754 1.805 1.753 1.770c
0.05 1.781 1.770 1.779 1.756 1.772c
0.10 1.799 1.764 1.764 1.767 1.773c
0.15 1.831 1.779 1.786 1.791 1.797b
0.20 1.869 1.838 1.828 1.826 1.840a
Average 1.809x 1.781yz 1.792y 1.779z
0–56 d feed:gain ratio
0.00 1.904 1.874 1.938 1.849 1.891c
0.05 1.904 1.890 1.891 1.881 1.891c
0.10 1.916 1.892 1.882 1.898 1.897c
0.15 1.945 1.898 1.919 1.912 1.919b
0.20 1.994 1.959 1.928 1.956 1.959a
Average 1.933x 1.903y 1.911y 1.899y
0–14 d 0–35 d 0–42 d 0–49 d 0–56 d
Source of
variance P diff SEM P diff SEM P diff SEM P diff SEM P diff SEM

Met level 0.41 0.008 0.0001 0.007 0.0001 0.007 0.0001 0.005 0.0001 0.008
Supplements 0.08 0.007 0.0002 0.006 0.0001 0.006 0.0001 0.005 0.004 0.007
Met × supplements 0.049 0.016 0.16 0.014 0.08 0.015 0.06 0.012 0.15 0.016
CV 2.79 1.90 1.78 1.24 1.72
Means within an age with common superscripts do not differ significantly (P < 0.05).
a–d

Means within a row with common superscripts do not differ significantly (P < 0.05).
x–z

fected in a linear manner. One of the first re-
ports suggesting that amino acid levels in ex-
cess of NRC recommendations improved breast
meat yield was that of Hickling et al. [31], who
fed broilers a combination of 2 Met levels (100
or 116% of 1984 NRC) with 4 levels of Lys
(100, 106, 112, or 118 of 1984 NRC). The
study by Hickling et al. [31] indicated that Lys
above NRC levels did not improve breast meat
yield unless Met was also greater than sug-
gested by NRC. However, only 2 Met levels
were compared (NRC, and 116%). Moran [32]
WALDROUP ET AL.: INDUSTRY BROILER DIETS
Figure 1. Feed conversion at different ages as influence by removal of various reductions in level of Met in the diet.
fed 2 strains of broilers (Ross × Ross; Steggles
× Arbor Acres) diets either deficient or ade-
quate in Met from 0 to 8 wk with processing
at both 6 and 8 wk. The low Met diets reduced
BW and breast yield and increased abdominal
fat at 6 wk but not at 8 wk of age. The deficient
diets fed from 6 to 8 wk were very near to the
NRC [30] requirement. Wallis [33] demon-
strated a response in breast meat yield to Met
supplementation of a deficient diet but did not
make any estimates about requirements for
maximum yield.
One of the few reports that attempt to titrate
a Met requirement to optimize breast yield was
that of Schutte and Pack [34]. Feeding a range
of Met or TSAA levels from 14 to 38 d, they
estimated a TSAA requirement of 0.84% for
BW gain, 0.88% for FCR, and 0.89% for breast
yield. It is difficult to compare the recommen-
65
dations of Schutte and Pack [34] to NRC [30] as
different time periods are involved. In a recent
study from our laboratory using NRC [30] feed-
ing intervals, Si et al. [35] noted that increasing
Met above NRC recommendations improved
FCR at 42 and 56 d but had no effect on breast
meat yield.
The ability of BET to spare some of the Met
needs in broiler diets is subject to considerable
controversy. Virtanen and Rosi [36] conducted
studies in which the basal diet was formulated
to provide 75% of the NRC [30] recommenda-
tion for Met. This diet was then fortified with
Met at 0.05, 0.10, or 0.15% or with BET at the
same levels. There was a significant response
in BW to additions of Met and BET; the authors
concluded that the response to BET was greater
than that obtained from Met. Virtanen and
Rumsey [18] summarized a number of unpub-
66 JAPR: Research Report

Table 8. Effects of levels of methionine, choline, and betaine on mortality of male broilers (means of 4 pens of
60 males/treatment)

500 mg
of choline/kg
Reduction No choline 1,000 mg 1,000 mg 500 mg
in Met (%) or betaine of choline/kg of betaine/kg of betaine/kg Average

0–14 d mortality (%)

0.00 4.16 2.91 2.91 2.70 3.17
0.05 2.08 5.00 3.33 4.37 3.69
0.10 2.08 5.55 2.91 3.33 3.47
0.15 4.58 6.25 3.33 3.75 4.47
0.20 3.33 2.91 2.50 2.91 2.91
Average 3.25 4.52 3.00 3.41
0–35 d mortality (%)
0.00 5.83 3.33 3.33 4.79 4.32
0.05 2.50 6.25 3.75 5.20 4.42
0.10 3.75 6.66 6.25 4.79 5.36
0.15 6.25 8.33 4.16 5.20 5.98
0.20 5.00 3.75 2.50 4.58 3.95
Average 4.66 5.66 4.00 4.91
0–42 d mortality (%)
0.00 5.83 4.58 4.58 5.20 5.05
0.05 5.41 7.50 3.75 5.83 5.62
0.10 4.16 8.33 7.08 5.83 6.35
0.15 8.33 9.16 5.41 6.25 7.29
0.20 6.25 4.58 2.91 7.08 5.20
Average 6.00 6.83 4.75 6.04
0–49 d mortality (%)
0.00 6.66 6.66 5.00 6.87 6.30
0.05 6.25 8.75 4.16 6.66 6.45
0.10 5.00 10.00 9.58 6.66 7.81
0.15 8.75 9.58 6.66 8.12 8.28
0.20 9.58 4.58 3.75 10.83 7.18
Average 7.25 7.91 5.83 7.83
0–56 d mortality (%)
0.00 9.58 8.33 5.83 8.33 8.02
0.05 7.91 11.66 5.41 7.29 8.07
0.10 9.16 12.77 10.41 9.16 10.38
0.15 10.83 13.75 8.75 9.79 10.78
0.20 12.50 7.50 4.58 12.91 9.37
Average 10.00 10.80 7.00 9.50
0–14 d 0–35 d 0–42 d 0–49 d 0–56 d
Source of
variance P diff SEM P diff SEM P diff SEM P diff SEM P diff SEM

Met level 0.54 0.66 0.37 0.79 0.31 0.83 0.56 0.98 0.33 1.17
Supplements 0.35 0.63 0.49 0.75 0.34 0.79 0.34 0.93 0.10 1.12
Met × supplements 0.91 1.41 0.70 1.69 0.74 1.78 0.34 2.09 0.64 2.51
CV 80.33 70.09 60.90 58.64 54.87

lished research trials conducted at private re-
search institutions using corn-soybean diets
with various levels of coccidiosis challenge.
They concluded that under practical field con-
ditions with chicks maintained on built-up litter
that BET was more effective in promoting
growth and feed efficiency than Met. However,
Rostagno and Pack [15], using corn-sorghum-
soybean meal diets supplemented with CHO to
provide methyl groups, found small and non-
significant responses to BET supplementation
that were not comparable to those provided by
supplemental Met. Schutte et al. [16] fed a
complex diet (corn, wheat, tapioca, peas,
WALDROUP ET AL.: INDUSTRY BROILER DIETS 67

Table 9. Effects of levels of methionine, choline, and betaine on dressing percentage of male broilers at different
ages (means of 4 groups of 5 birds/treatment)

500 mg
of choline/kg
Reduction No choline 1,000 mg 1,000 mg 500 mg
in Met (%) or betaine of choline/kg of betaine/kg of betaine/kg Average

42 d dressing percentage (%)

0.00 70.17 70.26 69.70 70.65 70.20
0.05 69.81 71.27 71.19 70.14 70.60
0.10 70.49 72.48 71.47 72.54 71.75
0.15 70.93 70.74 71.39 70.82 70.97
0.20 70.37 71.09 71.32 71.50 71.07
Average 70.35 71.17 71.01 71.13
49 d dressing percentage (%)
0.00 70.68 72.75 72.44 72.11 72.00
0.05 71.62 71.41 73.11 72.94 72.27
0.10 72.26 72.85 72.51 72.20 72.46
0.15 73.03 73.25 72.84 72.61 72.93
0.20 71.94 72.12 73.35 72.67 72.52
Average 71.91 72.48 72.85 72.51
56 d dressing percentage (%)
0.00 71.65 72.75 73.16 73.02 72.64
0.05 71.82 73.27 73.56 73.51 73.04
0.10 72.99 72.58 73.50 73.59 73.17
0.15 73.84 73.47 73.73 73.01 73.51
0.20 73.47 73.16 74.11 72.03 73.19
Average 72.45 73.04 73.61 73.03
42 d 49 d 56 d
Source of
variance P diff SEM P diff SEM P diff SEM

Met level 0.08 0.38 0.34 0.32 0.33 0.28

Supplements 0.31 0.34 0.15 0.28 0.13 0.25
Met × supplements 0.89 0.77 0.59 0.64 0.30 0.57
CV 2.18 1.76 1.58

feather meal, and soybean meal) and a simple
corn-soybean meal diet, both supplemented
with CHO and considered to be Met deficient,
and evaluated the effects of supplemental BET.
The response of chicks to BET was small and
nonsignificant, although BET significantly im-
proved oven-ready yield and breast meat yield.
However, the effect on breast yield was inferior
to that obtained by Met supplementation.
Esteve-Garcia and Mack [19] evaluated the
potential replacement value of Met by BET in
a Met-deficient diet. Choline was added at 500
mg/kg as a methyl donor, and clean conditions
were used to reduce coccidiosis challenge. There
were no significant interactions between BET
and Met; Met supplementation significantly im-
proved BW and feed to gain at 21 and 41 d
and significantly increased breast yield at 41 d.
Addition of BET significantly improved carcass
yield (dressing percentage) with no significant
change in other carcass parameters such as breast
yield or abdominal fat. Esteve-Garcia and Mack
[19] concluded that BET does not replace Met
in its function as an essential amino acid in
protein metabolism but may improve carcass
yield. McDevitt et al. [20] also noted that addi-
tion of BET to diets containing added Met fur-
ther improved the relative breast muscle yield.
They concluded that BET could not substitute
for Met in diets marginally deficient in TSAA
but that BET may improve carcass composition,
especially breast meat yield. Remus [37] noted
that breast meat yield of tom turkeys was sig-
nificantly improved by BET supplementation.
The results of the present study indicate that
addition of either CHO or BET at 1,000 mg/kg
to a nutritionally adequate corn-soybean meal
diet had little or no sparing effect on Met. Addi-
68 JAPR: Research Report

Table 10. Effects of levels of Met, choline, and betaine on breast yield at different ages (means of 4 groups of 5
birds/treatment)

500 mg
of choline/kg
Reduction No choline 1,000 mg 1,000 mg 500 mg
in Met (%) or betaine of choline/kg of betaine/kg of betaine/kg Average
1
42 d breast yield (%)
0.00 24.76 24.48 25.20 24.62 24.76a
0.05 24.27 25.19 24.54 24.83 24.71a
0.10 23.69 24.73 24.29 24.31 24.25ab
0.15 22.42 24.15 24.76 24.18 23.88b
0.20 21.83 23.46 23.59 23.60 23.12c
Average 23.39y 24.40x 24.48x 24.31x
49 d breast yield (%)
0.00 25.11 26.55 25.34 25.94 25.73a
0.05 24.33 25.29 25.97 25.70 25.32ab
0.10 24.37 24.97 23.94 25.78 24.76bc
.15 23.45 25.13 24.91 24.12 24.40c
0.20 21.54 23.30 23.66 23.56 23.01d
Average 23.76y 25.05x 24.76x 25.02x
56 d breast yield (%)
0.00 25.55 24.86 26.20 26.15 25.69a
0.05 25.23 25.29 25.86 25.34 25.43ab
0.10 24.01 25.07 25.55 26.13 25.19ab
0.15 24.49 25.07 25.42 25.27 25.06b
0.20 22.60 24.61 24.76 23.69 23.92c
Average 24.38z 24.98y 25.56x 25.32xy
42 d 49 d 56 d
Source of
variance P diff SEM P diff SEM P diff SEM

Met level 0.0005 0.27 0.0001 0.22 0.0001 0.22

Supplements 0.01 0.25 0.0001 0.19 0.0005 0.19
Met × supplements 0.70 0.55 0.08 0.43 0.16 0.43
CV 4.62 3.52 3.50

Means within an age with common superscripts do not differ significantly (P < 0.05).
a–d

Means within a row with common superscripts do not differ significantly (P < 0.05).
x–z
1
Skinless, boneless breast meat (pectoralis major + pectoralis minor) as a percentage of carcass weight.

tion of CHO or BET to the diets did improve
feed conversion at 35 or 42 d but not at other ages
evaluated. One of the most significant responses
observed during the present study was the posi-
tive influence of CHO or BET supplementation
on breast meat yield. In agreement with that of
many previous reports, addition of 1,000 mg of
BET/kg resulted in increased breast yield that
was independent of the Met content of the diet.
However, similar improvements were noted with
the addition of 1,000 mg of CHO/kg or a combi-
nation of BET and CHO, each at 500 mg/kg.
Due in large measure to its osmoregulatory
functions [17, 38, 39], BET appears to be of
benefit in chicks infected with coccidiosis [39,
40, 41, 42, 43, 44]. Chicks in the present study
received an anticoccidial in the diet and were
grown on new litter with no apparent problems
with coccidial infection. Intestinal strength of
chicks in the present study was not influenced by
supplementation of the diet with BET or CHO.
The Arg levels in this study exceeded those
suggested by NRC [30] but were similar to levels
used in the commercial broiler industry [21].
Keshavarz and Fuller [45, 46] demonstrated that
high levels of Arg in a corn-soybean meal diet
suppressed performance when Met levels were
low and suggested that the adverse effect of
excess Arg is due to the increased demand for
methyl groups for the formation of creatine,
which in turn leads to a deficiency of Met. This
research was confirmed by the findings of Cham-
WALDROUP ET AL.: INDUSTRY BROILER DIETS 69

Table 11. Effects of levels of methionine, choline, and betaine on tensile strength of intestine of male broilers
(means of 4 groups of 5 birds/treatment)

500 mg
of choline/kg
Reduction No choline 1,000 mg 1,000 mg 500 mg
in Met (%) or betaine of choline/kg of betaine/kg of betaine/kg Average

42 d peak load (kg)

0.00 0.412 0.403 0.443 0.454 0.428
0.05 0.424 0.530 0.361 0.466 0.445
0.10 0.508 0.461 0.505 0.404 0.469
0.15 0.413 0.387 0.473 0.395 0.417
0.20 0.408 0.432 0.418 0.436 0.423
Average 0.433 0.442 0.440 0.431
49 d peak load (kg)
0.00 0.558 0.760 0.492 0.549 0.590
0.05 0.654 0.594 0.528 0.553 0.582
0.10 0.543 0.498 0.620 0.578 0.560
0.15 0.510 0.523 0.543 0.492 0.517
0.20 0.567 0.530 0.547 0.529 0.543
Average 0.566 0.581 0.546 0.540
56 d peak load (kg)
0.00 0.542 0.457 0.552 0.600 0.538
0.05 0.490 0.538 0.600 0.531 0.540
0.10 0.589 0.498 0.543 0.552 0.545
0.15 0.475 0.529 0.521 0.588 0.528
0.20 0.535 0.557 0.530 0.513 0.534
Average 0.526 0.516 0.549 0.557
42 d 49 d 56 d
Source of
variance P diff SEM P diff SEM P diff SEM

Met level 0.49 0.023 0.41 0.029 0.99 0.025

Supplements 0.97 0.020 0.65 0.027 0.52 0.024
Met × supplements 0.34 0.046 0.26 0.061 0.73 0.050
CV 21.29 22.03 18.77

ruspollert et al. [47, 48]. Arg levels used in the increased demand for methyl groups a greater
present study were not as high as those shown response to BET or CHO should have been ob-
in the cited studies, and if there had been an served.

CONCLUSIONS AND APPLICATIONS

1. Dietary Met levels typical of top industry producers could be reduced by approximately 0.10%
without adversely affecting BW gain or feed conversion.
2. Reductions in Met levels below those used by top industry producers had a significant negative
effect on breast yield.
3. Addition of CHO or BET at 1,000 mg/kg or a combination of 500 mg/kg of both appeared to
have no sparing effect on Met needs.
4. Addition of CHO or BET at 1,000 mg/kg or a combination of 500 mg/kg of both significantly
improved feed conversion and breast meat yield of broilers independent of dietary Met levels.
5. No influence of CHO or BET supplementation was observed on intestinal strength of broilers
in the present study in which coccidiosis was not an apparent problem.
70
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Acknowledgments
The authors express their appreciation to Bill Huff and Susan
Watkins for their contributions as reviewers. Thanks are given to
Degussa Corporation for amino acid analysis of the diets and to
Danisco Animal Nutrition for contributing the betaine and for choline
and betaine analysis of the diets.