METABOLISM AND NUTRITION

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1 METABOLISM AND NUTRITION Effects of Reducing Dietary Protein, Methionine, Choline, Folic Acid, and Vitamin B 12 During the Late Stages of the Egg Production Cycle on Performance and Eggshell Quality 1 K. Keshavarz 2 Department of Animal Science, Cornell University, Ithaca, New York ABSTRACT A series of four experiments was conducted with no adverse effect on egg production. In experiment to determine whether shell quality during the late stages of egg production can be improved by using diets that are effective in reducing egg size. The experiments involved dietary manipulation of protein, methionine, choline, folic acid, and vitamin B 12. In experiment 1, reducing dietary protein in combination of reducing the dietary methionine and choline or this diet without supplemental folic acid and vitamin B 12 resulted in reduced egg weight and improved shell quality. However, egg production also was drastically reduced. In experiment 2, reducing the dietary level of methionine, without adding supplemental choline, folic acid, and vitamin B 12 reduced egg size and improved shell quality, but egg production was reduced as well. In this experiment reducing the dietary methionine without supplemental folic acid and vitamin B 12 reduced egg size and improved shell quality 3, reducing the dietary level of methionine and choline or reducing the dietary level of choline, folic acid, and vitamin B 12 reduced egg size and improved shell quality without adverse effects on egg production. On the other hand, reducing dietary methionine, folic acid, vitamin B 12, and supplemental choline reduced egg weight and improved shell quality but lowered egg production. In experiment 4, reducing dietary methionine together with reducing choline and vitamin B 12 reduced egg size and improved shell quality with no adverse effect on egg production. The results of this series of experiments generally indicate that certain manipulations of the combination of methionine, choline, folic acid, and vitamin B 12 have the potential to reduce egg weight and improve shell quality without affecting egg production during the latter stages of the egg production cycle. (Key words: late egg size, methionine, choline, folic acid, vitamin B 12 ) 2003 Poultry Science 82: INTRODUCTION It is estimated that 6 to 8% of eggs produced are lost in different phases of egg handling systems (Hamilton et al., 1979). Roland (1988) estimated that the loss to egg industry is about $480 million annually in the U.S. Shell quality problems become particularly serious during the high environmental temperatures of summer months and during the late stages of the egg production cycle. The results of a study by Roland et al. (1975) and data generated in our laboratory (Keshavarz and Nakajima, 1993) indicate that reduced shell quality with aging is not due to a reduced ability of hens to absorb or mobilize Ca for shell formation. Absolute daily retention of Ca in our experiment, and shell weight in Roland s experiment (1975), remained constant as hens aged. The reason for 2003 Poultry Science Association, Inc. Received for publication September 13, Accepted for publication April 17, Supported, in part, by ISA Babcock, P.O. Box 280, Ithaca, NY To whom correspondence should be addressed: kk33@cornell.edu. reduced shell quality is due to increased egg size, distributing a constant amount of shell over a larger egg. Consequently, limiting egg size should also prevent loss of shell thickness. Ousterhout (1980) reported that increasing dietary Ca with age resulted in improvement of shell quality due to limiting the increase in egg weight. Jackson et al. (1987) observed that egg weight was reduced and shell strength increased by reducing dietary methionine. However, these changes were obtained at the expense of lower egg production. Additionally, reducing the dietary nonphytate P (NPP) from 0.65 to 0.25% under the conditions of that experiment resulted in a reduction of egg weight and improvement in shell strength without affecting egg production. Petersen et al. (1983) reported that reducing dietary methionine reduced egg weight and improved shell quality without affecting egg production. Roland (1980), from an experiment with old hens, concluded that reducing egg size by dietary manipulation of protein, amino acids, energy, and Ca also reduced shell deposition and thus prevented any improvement in shell quality. We were not able to increase shell quality during the late stages of the egg production cycle by a number of dietary means, including increasing 1407

2 1408 KESHAVARZ the dietary Ca, reducing the dietary phosphorus, increasing the dietary level of vitamin D 3, or a combination of these factors (Keshavarz and Nakajima, 1993). However, shell quality of old hens was increased when 50% of supplemental Ca was provided by oyster shell. Increasing the concentration of several nutrients in the diet increases egg weight. These nutrients include methionine (Leong and McGinnis, 1952), choline (Tsiagbe et al., 1982; Keshavarz and Austic, 1985; Miles et al., 1986), and vitamin B 12 (Skinner, 1951; Squires and Naber, 1992). Combinations of related nutrients were methionine, choline, vitamin B 12 (Griffith et al., 1969), and homocystine and choline (Welch and Couch, 1955). The objective of the research reported here was to feed the minimum amounts of the above nutrients needed to maintain egg production but reduce egg weight. To the best of our knowledge, the effects on late egg size from dietary manipulation of methionine and methyl group contributions (choline, folic acid, and vitamin B 12 ) have not been investigated in a single experiment. Experiment 1 MATERIALS AND METHODS Three hundred seventy-five, 54-wk-old hens (Babcock B300) were used in the experiment. The birds were housed in groups of five birds per cage ( cm) in a windowless laying house. The birds of three adjacent cages were considered an experimental replicate, and five such replicates (75 hens/treatment) were fed each dietary treatment from 54 to 72 wk of age. The following treatments (Table 1) were prepared: T 1 = 16.0% protein to meet NRC (1994) requirements with added choline, folic acid, and vitamin B 12. T 2 = same as T 1 except with 13% protein. T 3 = same as T 2 except methionine deletion. T 4 = same as T 3 except without choline supplement. T 5 = same as T 4 except without folic acid and vitamin B 12 supplements. For formulation of the diets, choline, folic acid, and vitamin B 12 were removed from the vitamin mixture, and appropriate levels of these nutrients were added to the diets according to the design of the experiment. The diets were isocaloric, and with the exception of T 1 were isonitrogenous. Based on BW at the start of the experiment, 2 wk preexperimental egg production, and 3 consecutive d preexperimental egg weight records, means of these traits for different treatments were kept similar at the start of the experiment (P > 0.05). Records of daily egg production and weekly feed consumption were kept throughout the experiment and were summarized biweekly. All eggs produced during the last 4 d of the biweekly period were used for measurements of egg weight, egg grades (according to USDA grading system), and specific gravity. Specific gravity was determined using salt solutions varying in specific gravity from to in increments of units. BW was determined at the end of the experiment. Experiment 2 Two hundred eighty Babcock B300 hens were used in this experiment. The birds were housed in individual cages (30 45 cm). The birds of five adjacent cages were considered an experimental replicate, and each dietary treatment was fed to seven replicates (35 birds/treatment) for 12 wk (56 to 68 wk of age). Similar to the procedure used in experiment 1, average BW, egg production, and egg weight were not different among various treatment groups (P > 0.05) at the start of the experiment. The following treatments (Table 1) were prepared: T 1 = 15% protein to meet NRC (1994) requirements with added choline, folic acid, and vitamin B 12. T 2 = same as T 1 except 0.05% less methionine. T 3 = same as T 1 except for choline deletion. T 4 = same as T 1 except for folic acid and vitamin B 12 deletion. T 5 = combination of T 2 and T 3. T 6 = combination of T 2 and T 4. T 7 = combination of T 3 and T 4. T 8 = combination of T 2,T 3, and T 4. Records of daily egg production and weekly feed consumption were kept during the experiment and were summarized every 4 wk. All the eggs laid on 3 consecutive d on a biweekly basis were saved for measurements of egg weight, grade, and specific gravity. The procedure for measurement of specific gravity was similar to that described for experiment 1. Experiment 3 Six hundred 60-wk-old hens (Babcock B300) were used in this experiment. The birds were housed in groups of five birds per shallow cage (61 36 cm) in the same windowless house used in experiment 1. The birds of four adjacent cages were considered an experimental replicate, and each dietary treatment was fed to five such replicates (100 hens/treatment) during the experiment (60 to 72 wk of age). Similar to procedures used in previous experiments, the treatment means for BW, egg production, and egg weight were kept similar at the start of the experiment (P > 0.05). Corn was 68.55% and soybean meal was 20.86% of the diet used in this experiment. The following treatments (Table 1) were prepared: T 1 = 15% protein to meet NRC (1994) requirements with added choline, folic acid, and vitamin B 12. T 2 = same as T 1 except 0.05% less methionine and choline deletion. T 3 = same as T 1 with deletion of choline, folic acid, and vitamin B 12. T 4 = same as T 1 except 0.05% less methionine and deleted folic acid and vitamin B 12.

3 REDUCING EGG WEIGHT DURING THE LATE EGG PRODUCTION 1409 TABLE 1. Composition of experimental diets Ingredient 16% CP 13% CP 15% CP Corn Barley Soybean meal, dehulled Lysine HCl, 98% 0.08 Limestone Mono-dicalcium phosphate Salt Vitamin mixture Mineral mixture DL-Methionine Choline chloride, 3 60% Folic acid, 3 98% Vitamin B Calculated analysis 4 Energy (kcal ME/kg) 2,801 2,801 2,851 Protein (%) Lysine (%) Methionine (%) TSAA (%) Choline 5 (mg/kg) Total choline (mg/kg) 1,256 1,256 1,135 Folic acid 6 (mg/kg) Total folic acid (mg/kg) Vitamin B 12 (µg/kg) Linoleic acid (%) Calcium (%) Nonphytate P (%) Vitamin premix provided per kilogram of diet: vitamin A (retinyl acetate), 8,800 IU; cholecalciferol, 2,200 IU; DL-α-tocopheryl acetate, 11 IU; menadione sodium bisulfite, 2.2 mg; riboflavin, 4.4 mg; D-calcium pantothenate, 8.8 mg; nicotinic acid, 44 mg; pyridoxine hydrochloride, 2.2 mg; d-biotin, 0.11 mg; thiamine hydrochloride, 2.5 mg; ethoxyquin, 125 mg. 2 Mineral premix provided per kilogram of diet: MnSO 4 H 2 O, 185 mg; ZnO, 62 mg; FeSO 4 7H 2 O, 149 mg; CuSO 4 5H 2 O, 19.6 mg; KI, 1.4 mg; Na 2 SeO 3, 0.22 mg. 3 Appropriate levels of choline chloride 60%, folic acid 98%, and vitamin B 12 were added to complete appropriate diets. 4 Based on tables of feed composition of NRC (1994) except that the determined protein contents of corn (7.4%), barley (11.5%), and soybean meal (48.4%) were used in formulating the experimental diets. During wk 15 to 18, a new shipment of barley with determined protein content of 9% was used. By slightly increasing the level of soybean meal and reducing the level of barley, the protein level was maintained at 13% in T 2 to T 5 for the period of 15 to 18 wk of the experiment. 5 From choline chloride, 60%. 6 From folic acid, 98%. (%) T 5 = same as T 1 except 0.05% less methionine and deleted choline, folic acid and vitamin B 12. T 6 = same as T 5 except 0.03% less methionine. New shipments of corn and soybeans with determined protein and dry matter contents of 7.59 and 87.8%, respectively, for corn and 46.8 and 89.5%, respectively, for soybean meal were used in this experiment. By using determined protein contents of corn and soybeans and the regression equations suggested by NRC (1994), the lysine, methionine, and TSAA contents of the corn and soybeans were calculated and adjusted for their dry matter content. The new amino acids values obtained were used in formulating the experimental diets. The daily feed consumption was intentionally maintained at about 100 g/hen per d during the experiment to have better control of the daily nutrient intake. An extra replicate for each dietary treatment was maintained in this experiment. When a bird from the experimental groups died, it was replaced with a random sample bird from the extra replicate, which was consuming a similar diet. This approach facilitated the daily feed delivery of about 100 g/hen per d. Similar to previous experiments, records of daily egg production and weekly feed consumption were kept during the experiment. All eggs laid on 3 consecutive d and on a biweekly basis were saved for measurements of egg weight, grade, and specific gravity. Salt solutions used for the measurement of specific gravity were identical to those used in previous experiments. Body weight was determined at the end of the experiment (72 wk of age). Experiment 4 Four hundred eighty Babcock B wk-old hens were used in this experiment. Four replicates of 15 hens were fed each dietary treatment for 10 wk (62 to 72 wk of age). The birds of each replicate were housed in groups of five in three adjacent shallow cages. Similar to previous experiments based on 2-wk pre-experimen-

4 1410 KESHAVARZ tal egg production, 3 d pre-experimental egg weight, and BW at the start of the experiment, the treatment means were kept similar (P > 0.05) at the start of the experiment. Records of daily egg production and weekly feed consumption were kept during the experiment. BW was determined at the end of the experiment. All the eggs produced during the last4dofevery 28-day period were used for measurements of egg weight, grade, and specific gravity. The basal diet used in this experiment contained 70.11% corn and 19.3% soybean meal. Experimental diets were formulated based on determined protein and amino acid content of corn and determined protein of soybean meal. Also, from the protein content of soybean meal and using the regression equations suggested by NRC (1994) to estimate methionine, TSAA, and lysine from the protein content of soybean meal, these amino acids were estimated and these values were used in feed formulation. The following treatments (Table 1) were prepared: T 1 = 15% protein to meet the NRC (1994) requirements with added choline, folic acid and vitamin B 12. T 2 = same as T 1 except 0.05% less methionine and choline deletion. T 3 = same as T 1 except 0.05% less methionine and deletion of vitamin B 12. T 4 = same as T 1 except 0.05% less methionine and deletion of folic acid. T 5 = same as T 1 with deletion of choline and vitamin B 12. T 6 = same as T 1 with deletion of choline and folic acid. T 7 = same as T 2 with deletion of vitamin B 12. T 8 = same as T 2 with deletion of folic acid. The data from each experiment were analyzed by AN- OVA of SAS software (2001), and means were compared by Duncan s multiple-range test (1955). Significance was considered as P All experiments were conducted according to an institutional animal care and use committee approved protocol in a facility accredited by the Association for Assessment and Accreditation of Laboratory Animal Care. Experiment 1 RESULTS AND DISCUSSION The results of experiment 1 are shown in Table 2. None of the production traits was decreased when protein level was reduced from 16 to 13%, but the levels of supplemental methionine, choline, folic acid, and vitamin B 12 were equal to the control diet (T 1 vs. T 2 ). When dietary level of protein was reduced to 13% and supplemental methionine was omitted from the diet (T 3 ), egg production, egg weight, egg mass, feed consumption, BW gain, and extra-large plus large-sized eggs were reduced, and feed conversion and small plus peewee-sized eggs were increased (T 1 vs. T 3 ). Specific gravity was TABLE 2. The effect of dietary manipulation of nutrients on production performance and specific gravity (54 to 72 wk, experiment 1) Egg grade Body Egg Egg Egg Feed Feed weight Large Small and production weight mass consumption conversion change and above Medium peewee Specific Treatment (%) (g) (g) (g) (g:g) (g) (%) (%) (%) gravity T 1, 16% protein (control) 73.9 a 58.2 a 43 a 98.1 a 2.29 c 53 a 47.4 a 50.2 a 2.3 c cd 2 T, 13% protein 71.1 a 57.2 a 40.7 a 93.9 ab 2.31 c 79 a 37.6 a 58.2 a 4.1 c d 3 2 T,asT minus methionine 56.8 b 54.6 b 31 b 92.2 bc 2.99 b 153 b 22.2 b 61.3 a 16.5 b bc 4 2 T,asT minus methionine and choline 46.3 c 53.1 b 24.6 c 86.2 c 3.56 a 153 b 16.7 b 56.6 a 26.7 b b T 5,asT 2 minus methionine, choline, 12 folic acid and vitamin B 48.2 c 51.2 c 24.7 c 88.0 c 3.59 a 212 b 9.5 b 51.2 a 39.2 a a SEM a d Means followed by different superscripts in each column are significantly different (P < 0.05).

5 REDUCING EGG WEIGHT DURING THE LATE EGG PRODUCTION 1411 slightly, but not significantly greater for hens fed a diet without supplemental methionine than the control group (T 1 vs. T 3 ). When in addition to supplemental methionine, supplemental choline (T 4 ) or supplemental choline and supplemental folic acid and vitamin B 12 (T 5 ) were also deleted from the 13% protein diet, production traits were reduced further and specific gravity was increased stepwise. The results of experiment 1 indicated that it is possible to reduce egg weight and, as a result, increase shell quality by proper dietary manipulation of nutrients. However, the extent of reducing the dietary nutrients in this experiment was too severe and resulted in reduced egg production. Removal of folic acid and vitamin B 12 from the diet (T 5 ) did not produce any additional adverse effects on performance except that egg weight was reduced considerably and, due to this, shell quality was further increased. Experiment 2 Egg production was not different among dietary treatments (Table 3). The only exception was egg production of birds of T 8, which were fed a diet similar to the control group but lower in methionine and not supplemented with choline, folic acid, and vitamin B 12 (T 1 vs. T 8 ). With the exception of birds fed the control diet not supplemented with choline (T 3 ), egg weight of other groups was consistently lower than the control group. However, egg weight was significantly lower than the control group only for hens of T 6, which were fed the lower methionine diet not supplemented with folic acid and vitamin B 12, and the hens of T 8, which were fed a diet similar to T 6 and that also was not supplemented with choline (T 1 vs. T 6 and T 8 ). Egg mass and BW gain were lower than the control group only for hens fed T 8. Feed consumption and feed conversion were not different for birds fed various dietary regimens than those fed the control diet. The lower egg weights of birds fed T 6 and T 8 than the controls was reflected in production of fewer large and extra-large eggs and more medium and small plus peewee than the control group, although none of these differences were significant. Specific gravity was not influenced by reducing the dietary methionine or removing supplemental choline from the control diet (T 1 vs. T 2 and T 3 ). However, there was a tendency for improvement of specific gravity of eggs from hens fed the control diet not supplemented with folic acid and vitamin B 12 (T 4 ) and for hens fed the low methionine and not supplemented with choline (T 5 ) when compared to the control group eggs. Specific gravities of eggs produced by hens fed T 6 to T 8 were greater than those fed the control diet. Improvement in specific gravity, for the most part, was closely related to reduced egg weight attributed to use of different dietary regimens. The information from this experiment indicated that it is possible to reduce egg weight and increase specific gravity without affecting egg production (T 1 vs. T 6 ). TABLE 3. The effect of dietary manipulation of nutrients on production performance and shell quality (56 to 68 wk, experiment 2) Egg grades Body Egg Egg Egg Feed Feed weight Large Small and production weight mass consumption conversion change and above Medium peewee Specific Treatment (%) (g) (g) (g) (g:g) (g) (%) (%) (%) gravity T 1, 15% protein (control) 80.9 a 59.3 a 48.0 a a 2.18 ab 31 a 64 abc 35.7 ab 0.2 b b 2 1 T,asT minus methionine 79.2 ab 57.7 abc 45.9 a a 2.29 ab 62 ab 54.4 abcd 43.3 ab 2.3 ab b 3 1 T,asT minus choline 80.8 ab 59.3 a 47.9 a a 2.16 b 33 a 72.9 a 27.1 b 0.0 b b T 4,asT 1 minus folic acid 12 and vitamin B 79.0 ab 57.8 abc 45.6 a a 2.22 ab 64 ab 54.9 abcd 44.8 ab 0.3 b ab T 5,asT 1 minus methionine and choline 78.9 ab 57.2 abc 45.2 a a 2.29 ab 75 ab 52.2 bcd 42.6 ab 5.2 ab ab T 6,asT 1 minus methionine, folic 12 acid and vitamin B 79.8 ab 56.7 bc 45.3 a a 2.29 ab 68 ab 46.9 cd 52.3 a 0.8 b a T 7,asT 1 minus choline, folic 12 acid and vitamin B 78.7 ab 58.6 ab 46.1 a a 2.18 ab 104 ab 67 ab 31.4 b 1.5 ab a T 8,asT 1 minus methionine, choline 12 folic acid, and vitamin B 75.5 b 56.1 c 42.3 b 98.1 a 2.33 a 156 b 43.8 d 49.9 a 6.3 ab a SEM a d Means followed by different superscripts in each column are significantly different (P < 0.05).

6 1412 KESHAVARZ Experiment 3 Experiment 3 was conducted to gain additional information about the repeatability of the effect of T 5 to T 8 on egg weight and specific gravity. The T 2 to T 5 treatments in experiment 3 (Table 4) corresponded to T 5 to T 8 treatments in experiment 2. Additionally, because egg production was lower in experiment 2 for hens fed T 8 than the control groups (T 1 ), an additional treatment (T 6 ) was added in experiment 3. This treatment was similar to T 5 except that the supplemental methionine was reduced only by 0.03% in this diet to reduce the stress of methionine deficiency on egg production. Reducing dietary methionine together with deleting supplemental choline (T 2 ) reduced egg weight and extralarge plus large-sized eggs and increased specific gravity with no adverse effect on other production traits (Table 4). Removing supplemental choline and folic acid and vitamin B 12 from the control diet (T 3 ) had a similar effect on egg production, weight, grade, and specific gravity as reducing the dietary methionine and deleting choline from the control diet (T 1 vs. T 2 and T 3 ). The only exception was that egg mass also was reduced due to effect of T 3. Reducing dietary methionine plus deleting the supplemental folic acid and vitamin B 12 from the control diet (T 4 ), although resulting in reduced egg weight and extra-large and large eggs and improvement of specific gravity, had an adverse effect on egg production. When, in addition to reducing the dietary methionine and deleting the supplemental folic acid and vitamin B 12, supplemental choline was also removed from the control diet (T 5 ), egg weight and extra-large plus large eggs were reduced further, and a greater improvement in specific gravity was observed. However, egg production was reduced further, and feed consumption was reduced with this dietary treatment. The addition of 0.02% methionine to T 5 (i.e., increasing the TSAA from 0.53 to 0.55%) did not have any effect in improving the performance (T 5 vs. T 6 ). The results of experiment 3 (Table 4) indicated that reducing the dietary methionine and removing the supplemental choline from the diet (T 2 ) or removing supplemental choline, folic acid, and vitamin B 12 from the vitamin mix (T 3 ) had the potential to reduce egg size and improve shell quality without adversely affecting egg production. Experiment 4 The results of experiment 3, in combination with those of the previous experiments indicated that removal of supplemental folic acid and vitamin B 12 in addition to reducing the dietary level of methionine, or removal of choline, or their combination, decreased egg production. It was hypothesized that reducing the dietary level of methionine, or deleting the supplemental choline, or the combination of these, together with removing either the supplemental level of folic acid or vitamin B 12, may be less stressful on egg production but still would have the TABLE 4. The effect of dietary manipulation of nutrients on production performance and shell quality 1 (60 to 72 wk, experiment 3) Egg grade Body Egg Egg Egg Feed Feed weight Large Small and production weight mass consumption conversion change and above Medium peewee Specific Treatment (%) (g) (g) (g) (g:g) (g) (%) (%) (%) gravity T 1, 15% protein (control) 71.0 a 61.1 a 43.4 a a 2.35 c 117 a 74.5 a 24.5 a 0.9 b d 2 1 T,asT minus methionine and choline 71.3 a 59.3 b 42.2 ab ab 2.41 c 146 ab 64.2 b 33.8 b 2.1 b c T 3,asT 1 minus choline, folic 12 acid and vitamin B 67.2 ab 59.2 b 39.8 bc a 2.56 bc 187 bc 61.2 b 34.1 b 4.7 b b T 4,asT 1 minus methionine, folic 12 acid and vitamin B 64.4 bc 57.7 b 37.2 c abc 2.70 b 170 bc 45.9 c 48.7 a 5.4 b b T 5,asT 1 minus methionine, choline 12 folic acid and vitamin B 60.2 c 55.0 c 33.1 d 98.5 bc 2.99 a 186 bc 33.3 d 53.4 a 13.3 a a 6 5 T,asT plus 0.02% methionine 59.7 c 55.3 c 33.1 d 97.3 c 2.95 a 198 c 34.6 d 47.3 a 15.4 a ab SEM a d Means followed by different superscripts in each column are significantly different (P < 0.05). 1 TSAA was 0.58% in T1, 0.55% in T 6, and 0.53% in the other dietary treatments.

7 REDUCING EGG WEIGHT DURING THE LATE EGG PRODUCTION 1413 TABLE 5. The effect of dietary manipulation of nutrients on production performance and shell quality (62 to 72 wk, experiment 4) Egg grade Body Egg Egg Egg Feed Feed weight Large and Small and production weight mass consumption conversion change above Medium peewee Specific Treatment (%) (g) (g) (g) (g:g) (g) (%) (%) (%) gravity T 1, 15% protein (control) 70.7 ab 63.3 a 44.6 abc 97 b 2.17 ab 85 b 83.7 a 16.3 a b T 2,asT 1 minus methionine and choline 69.9 ab 61.4 ab 42.5 ab 98.5 ab 2.33 ab 116 ab 80.0 ab 19.8 a 0.2 a ab T 3,asT 1 minus methionine and vitamin B ab 61.9 ab 40.9 bc 97.9 ab 2.38 ab 103 ab 80.4 ab 19.3 a 0.3 a ab T 4,asT 1 minus methionine and folic acid 75.4 a 61.6 ab 45.9 ab 97.8 ab 2.11 b 166 a 72.9 ab 26.9 a 0.8 a ab T 5,asT 1 minus choline and vitamin B ab 62.6 ab 46.4 a 97.5 ab 2.08 b 126 ab 83.1 ab 16.5 a 0.4 a ab T 6,asT 1 minus choline and folic acid 67.3 ab 61.4 ab 41.0 bc 97.0 b 2.37 ab 116 ab 74.2 ab 25.6 a 0.3 a b T 7,asT 1 minus methionine, choline, and B b 60.4 b 39.9 c 96.9 b 2.43 a 108 ab 64.8 b 34.4 a 0.9 a a T 8,asT 1 minus methionine, choline, and folic acid 66.5 b 61.4 ab 40.5 c 96.6 b 2.39 ab 123 ab 74.0 ab 25.6 a 0.4 a ab SEM a c Means followed by different superscripts in each column are significantly different (P < 0.05). potential to reduce egg size and improve shell quality. Experiment 4 was conducted to test this hypothesis. As shown in Table 5, egg production of birds fed the control diet minus methionine and folic acid (T 4 ) was significantly greater than those fed the control diet minus methionine, choline, and vitamin B 12 (T 7 ) and those fed the control diet minus methionine, choline, and folic acid (T 8 ) (Table 5). Although egg weight was moderately lower for all treatments than the controls, only the differences between egg weight of the control group and the birds fed the control diet minus the combination of methionine, choline, and B 12 were different (T 1 vs. T 7 ). Egg mass, feed intake, feed conversion, and BW gain of birds fed different dietary treatments were not different from the controls. The exception was BW of birds fed the control diet minus the combination of methionine and folic acid (T 4 ) that was greater than the control group. Extra-large, medium, and small plus peewee eggs did not differ among the treatments. The exception was the extra-large plus large eggs of the birds fed the control diet minus the combination of methionine, choline, and B 12 (T 7 ), which was lower than the control group. Specific gravity was moderately greater for all the treatments than the control, although only the specific gravity of birds fed the control diet minus the combination of methionine, choline, and B 12 was greater than the control group (T 1 vs. T 7 ). To our knowledge there is no information in the literature regarding the effect of reducing the dietary level of choline, folic acid, and vitamin B 12 on egg weight, egg production, and shell quality of old hens that can be used for comparative purposes with those reported here. The information obtained from this series of experiments indicated that the potential exists for reducing egg weight and, in turn, increasing shell quality without loss of egg production, by certain manipulation of the combination of dietary level of methionine, choline, folic acid, and vitamin B 12. For example, in experiment 2 reducing the dietary methionine together with deleting supplemental folic acid and vitamin B 12 (T 6 ) reduced egg weight and improved shell quality with no adverse effects on egg production. Similarly, in experiment 3, reducing the dietary level of methionine and choline (T 2 ), or reducing the dietary level of choline, folic acid, and vitamin B 12 (T 3 ), reduced egg weight and improved shell quality without an adverse effect on egg production. Furthermore, in experiment 4, reducing dietary methionine together with reducing choline and vitamin B 12 (T 7 ), reduced egg weight and improved shell quality with no adverse effect on egg production. REFERENCES Duncan, D. B Multiple range and multiple F-tests. Biometrics 11:1 42. Griffith, M., A. J. Olinde, R. Schexnailder, R. F. Davenport, and W. F. McKnight Effect of choline, methionine, and vitamin B 12 on liver fat, egg production and egg weight in hens. Poult. Sci. 48: Hamilton, R. M. G., K. G. Hollands, P. W. Voisey, and A. A. Grunder Relationship between egg shell quality and shell breakage and factors that affect shell breakage in the field. A review. World s Poult. Sci. J. 35: Jackson, M. E., H. M. Hellwig, and W. P. Waldroup Shell quality: Potential for improvement by dietary means and relationship with egg size. Poult. Sci. 66: Keshavarz, K., and R. E. Austic An investigation concerning the possibility of replacing supplemental methionine with choline in practical laying rations. Poult. Sci. 64: Keshavarz, K., and S. Nakajima Re-evaluation of calcium and phosphorus requirement of laying hens for optimum performance and eggshell quality. Poult. Sci. 72: Leong, K. C., and J. McGinnis An estimate of methionine requirement for egg production. Poult. Sci. 31: Miles, R. D., N. Ruiz, and R. H. Harms Response of laying hens to choline when fed practical layer diets devoid of supplemental sulfur amino acids. Poult. Sci. 65:

8 1414 KESHAVARZ National Research Council Nutrient requirements of poultry. 9th rev. ed. National Academy Press, Washington, DC. Ousterhout, L. E Effect of calcium and phosphorus levels on egg weight and egg shell quality in laying hens. Poult. Sci. 59: Petersen, C. F., E. A. Sauter, E. E. Steele, and J. F. Parkinson Use of methionine intake restriction to improve egg shell quality by control of egg weight. Poult. Sci. 62: Roland, D. A., Sr., D. R. Sloan, and R. H. Harms The ability of hens to maintain calcium deposition in egg shell and egg yolk as the hen ages. Poult. Sci. 54: Roland, D. A. Sr Egg shell quality: I. Effects of dietary manipulations of protein, amino acids, energy and calcium in aged hens on egg weight, shell weight, shell quality and egg components. Poult. Sci. 59: Roland, D. A. Sr Research note: Egg shell problem: Estimate of incidence and economic impact. Poult. Sci. 67: SAS Institute Statistical Analysis System Proprietary Software. Release 8.2. SAS Institute Inc., Cary, NC. Skinner, J. L., J. H. Quisenberry, and J. R. Couch High efficiency APF concentrates in the ration of laying fowl. Poult. Sci. 30: Squires, W. M., and E. C. Naber Vitamin profiles of eggs as indicators of nutritional status in the laying hens: Vitamin B 12 study. Poult. Sci. 71: Tsiagbe, V. K., C. W. Kang, and M. L. Sunde The effect of choline supplementation in growing pullet and laying hen diets. Poult. Sci. 61: Welch, B. E., and J. R. Couch Homocystine vitamin B 12, choline, and methionine in the nutrition of laying fowl. Poult. Sci. 34:

Department of Animal Sciences, University of Florida, Gainesville, Florida 32611

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