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The Influence of Feeding Diets Containing Different Calcium-Phosphorus Ratios on the Laying Hen1
ARGININE-LYSINE IN RAPESEED
Sharby, T. F., and J. M. Bell, 1974. In: Research on Rapeseed Seed, Oil and Meal. 3rd Progress Rep. Res. Committee, Rapeseed Assoc, Canada. Steel, R. G. D., and J. H. Torrie, 1960. Principles and Procedures of Statistics. McGraw-Hill Book Company Inc., Toronto. Stutz, M. W., J. E. Savage and B. L. O'Dell, 1971. Relation of dietary cations to arginine-lysine antagonism and free amino acid patterns in chicks. J. Nutr. 101: 377-394. Tao, R., R. J. Belzile and G. J. Brisson, 1971. Amino acid digestibility of rapeseed meal fed to chickens: effect of fat and lysine supplementation. Can. J. Anim. Sci. 51: 705-709. Vapar.Z., and D. R. Clandinin, 1972. Effect of tannins in rapeseed meal on its nutritional value for chicks. Poultry Sci., 51: 222-228. Williams, R. T., 1959. Cited in: Nesheim, M. C , and Garlich, J. D. 1963. Studies on ornithine synthesis in relation to benzoic acid excretion in the domestic fowl. J. Nutr. 79: 311-317.
The Influence of Feeding Diets Containing Different Calcium-Phosphorus Ratios on the Laying Hen1 D . A. ROLAND, S R . AND R. H . HARMS
Department ofPoultry Science, Florida Agricultural Experiment Station, Gainesville, Florida 32611 (Received for publication June 25, 1975)
ABSTRACT Six experiments were conducted to determine the influence of feeding diets containing different calcium-phosphorus (Ca:P) ratios at different times on the laying hen. Feeding a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 1.50% P from 3 p.m. to 8 p.m. decreased serum calcium in three of four experiments and decreased specific gravity of eggs in four of four experiments, when compared to that of the birds fed the control diet which contained 0.75% P during both time periods. Specific gravity of eggs from hens fed a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 0.33% P from 3 p.m. to 8 p.m. was increased in three of four experiments. Although the differences are not significant, with one exception, the trend in the experiments indicate that the level of P in the diet and time of intake can influence shell quality and serum calcium. The dietary treatments had no significant influence on feed consumption, egg production or egg weight during any of these experiments. It was postulated that the adverse or beneficial effect on specific gravity of eggs and serum calcium as a result of feeding diets containing various Ca:P ratios was due to the influence on calcium absorption and/or bone resorption. POULTRY SCIENCE 55: 637-641, 1976
INTRODUCTION
T
H E calcium-phosphorus (Ca:P) ratio is one of the best known concepts in poultry nutrition. Although it is extremely important in chick diets, it is generally believed that 1. Florida Agr. Exp. Sta. Journal Series No. 5972.
the C a : P ratio is not very important in laying h e n diets. M a n y studies h a v e been reported in which varying the p h o s p h o r u s c o n t e n t of the diet from 0.3 to 0.7% had no significant influence on egg shell quality (Crowley et al., 1961; E v a n s et al., 1944; Gillis et al., 1953; Singsen et al, 1962; Walter a n d Aitken,
Downloaded from http://ps.oxfordjournals.org/ at Oakland University on May 31, 2015
97-120 Pergomon Press, New York. Jones, J. D., 1964. Lysine-arginine antagonism in the chick. J. Nutr. 84: 313-321. Kadirvel, R., G. V. N. Rayudu and P. Vohra, 1969. Excretion of metabolites of tannic acid by chickens with and without ceca. Poultry Sci. 48: 1511-1513. Leslie, A. J., and J. D. Summers, 1975. Amino acid balance of rapeseed meal. Poultry Sci. 54: 532-537. Nesheim, M. C , and J. D. Garlich, 1963. Studies on ornithine synthesis in relation to benzoic acid excretion in the domestic fowl. J. Nutr. 79: 311. O'Dell, B. L., and J. E. Savage, 1966. Arginine-lysine antagonism in the chick and its relationship of dietary cations. J. Nutr. 90: 364-370. Olomu, J. M , A. R. Robblee and D. R. Clandinin, 1974. Effects of processing and amino acid supplementation on the nutritive value of rapeseed for broilers. Poultry Sci. 53: 175-184. Rayudu, G. V. N., R. Kadirvel, P. Vohra and F. H. Kratzer, 1970. Toxicity of tannic acid and its metabolites for chickens. Poultry Sci. 49: 957-960.
637
638
D. A. ROLAND, SR. AND R. H. HARMS
In the following studies, diets containing different Ca:P ratios were fed to hens at different periods during the day in an effort to create the best environment for maximum mobilization, utilization and repletion of skeletal calcium. EXPERIMENTAL PROCEDURE Experiment 1. One-hundred-and-twenty Babcock B-300 hens (hatched March 10, 1972)
were divided into three groups of 40 birds each on August 28, 1973 and fed one of the following treatments: The hens in group 1 were fed a control diet containing 0.75% phosphorus (P) from 6 a.m. to 11 a.m. and from 3 p.m. to 8 p.m. Hens in group 2 were fed a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 0.33% P from 3 p.m. to 8 p.m. (low phosphorus diet). Hens in group 3 were fed a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 1.50% P from 3 p.m. to 8 p.m. (high phosphorus diet). All diets contained 3.00% calcium and were iso-nitrogenous and iso-caloric. Hens were not given access to feed from 11 a.m. to 3 p.m. or from 8 p.m. to 6 a.m. Feed consumed during each feeding period was determined daily throughout the seven-day experiment. Specific gravity and egg weight were determined on all eggs. Blood was taken by anterior heart puncture from one-half of all birds at 6 a.m. on the eighth day and serum calcium determined. Experiment 2. The procedure of Experiment 2 was identical to Experiment 1 except that Babcock B-300 hens were hatched on May 22, 1973 and the experiment was initiated on October 29, 1974 and lasted for 10 days. Blood was taken at 6 a.m. on the 11th day. Experiment 3. One-hundred Babcock B-300 hens (hatched May 22, 1972) producing eggs with low specific gravity were divided into two groups. The hens were fed either a control diet containing 0.75% P from 6 a.m. to 11 a.m. and from 3 p.m. to 8 p.m. or a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 0.33% P from 3 p.m. to 8 p.m. Hens were not given access to feed from 11 a.m. to 3 p.m. or from 8 p.m. to 6 a.m. All diets contained 3.00% calcium and were iso-nitrogenous and iso-caloric. Feed consumed during each feeding period was determined daily throughout the 10-day experiment. Specific gravity and egg weight
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1962). More recent studies have indicated that phosphorus may have a more important role in calcium metabolism in the laying hen than previously believed. Holcombe et al. (1974) reported that the hen's tolerance of phosphorus was influenced by the calcium level of the diet and that extremely high levels of phosphorus had an adverse effect on shell quality. They also found that hens have the ability to adjust phosphorus intake when offered a choice of diets containing different levels of phosphorus. This ability of hens to regulate phosphorus intake was just as distinct as their ability to regulate calcium intake (Holcombe et al., 1975). Several recent studies have also indicated that phosphorus has a pronounced effect on calcium absorption and bone resorption (Raisz and Nieman, 1969; Morrissey and Wasserman, 1971). Cuisinier-Gleizes et al. (1971) reported that low phosphorus diets resulted in an immediate increase in bone resorption. The utilization of skeletal calcium for egg shell formation is apparently necessary and most important during the late hours of the night when total calcium and percent calcium of the feed contents of the digestive system is at its lowest (Roland et al., 1973). This release of calcium from the skeletal system into the blood is accompanied by a release of phosphorus. The increased amount of recycled phosphorus coupled with dietary phosphorus in the blood at this time may not be the best environment for maximum stimulation of bone resorption which is necessary for egg shell formation.
C A : P RATIOS
were determined on all eggs. Blood was taken by anterior heart puncture from one-half of all birds at 6 a.m. on the 11th day. Experiment 4. The procedure of Experiment 4 was similar to Experiment 3 except that 50 hens producing eggs with a high specific gravity were used.
Experiment 6. The procedure of Experiment 6 was identical to Experiment 5 except that 50 hens producing eggs with high specific gravity were used. The hens were fed either a diet containing 0.75% P from 6 a.m. to 11 a.m. and from 3 p.m. to 8 a.m. or a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 1.50% P from 3 p.m. to 8 p.m. In all experiments serum calcium was determined using an atomic absorption spectrophotometer. All data were statistically analyzed by analysis of variance (Snedecor, 1956), and significant differences were determined by the multiple range test of Duncan (1955). RESULTS AND DISCUSSION In three of four experiments a decrease in serum calcium occurred in hens fed the diet containing 1.50% P from 3 p.m. to 8 p.m. (Tables 1 and 3). The decrease in serum calcium approached significance at the 0.05 level in Experiments 1, 2 and 5. Serum calcium of hens fed the diet containing 0.33% P from 3 p.m. to 8 p.m. in Experiments 1-4 was numerically reduced (Tables 1 and 2).
Specific gravity of eggs from hens fed the diet containing 1.50% P from 3 p.m. to 8 p.m. was decreased in four of four experiments (Tables 1 and 3). Specific gravity of eggs from hens fed the diet containing 0.33% P from 3 p.m. to 8 p.m. was increased in three of four experiments (Tables 1 and 2). Dietary treatment had no significant influence on feed consumption, egg production or egg weight during the experiments. The hens consumed an average of 53% more feed from 3 p.m. to 8 p.m. than from 6 a.m. to 11 a.m. Feeding hens a diet containing a 2:1 Ca:P ratio from 3 p.m. to 8 p.m. appeared to have an adverse effect on serum calcium and specific gravity of eggs. However, feeding hens a diet containing a 9:1 Ca:P ratio from 3 p.m. to 8 p.m. had a beneficial effect on specific gravity of eggs in three of four experiments. In Experiment 2, specific gravity of eggs from hens fed the 9:1 Ca:P ratio during the evening was significantly greater than that from hens fed the diet containing a 2:1 Ca:P ratio during the same period. The results of these studies suggest that the Ca:P ratio of the diet fed to hens can influence serum calcium and specific gravity of eggs. It is believed that the response of hens in these experiments to various Ca:P ratios was due to the influence of phosphorus on calcium metabolism. Several studies have indicated that phosphorus has a pronounced effect on calcium absorption and bone resorption (Raisz and Nieman, 1969; Morrissey and Wasserman, 1971) Cuisinier-Gleizes et al. (1971) reported that low phosphorus diets resulted in an immediate increase in bone resorption. Several workers have also shown that calcium in the skeletal system is important in the process of egg shell formation (Tyler, 1940a, b; Taylor and Moore, 1958; Taylor, 1970; Candlish, 1971). Large quantities of calcium were mobilized from the skeletal system and utilized when calcium was not available from the intestinal contents
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Experiment 5. The procedure of Experiment 5 was similar to Experiment 3 except that Experiment 5 was initiated on April 15, 1974. The hens producing eggs with poor egg shell quality were fed either a diet containing 0.75% P from 6 a.m. to 11 a.m. and from 3 p.m. to 8 p.m. or a diet containing 0.33% P from 3 p.m. to 8 p.m.
639
640
D. A. ROLAND, SR. AND R. H. HARMS
TABLE 1.—Serum calcium and egg specific gravity from hens fed various dietary treatments (Exps. 1 and 2) Serum calcium (mg. %) Egg specific gravity Treatment Exp. 1 Exp. 2 Exp. Exp. 2 0.75%P l 6a.m.-lla.m. 2 3 26.9a ± 1.2 27.0a ± 1.2 1.071a ± 0.0011 1.073ab ± 0.0010 0.75% P 3 p.m.-8 p.m. 0.75% P 6 a.m.-11 a.m. 26.8a ±1.0 26.6a ± 0.9 1.072a ± 0.0009 1.074b ±0.0009 0.33% P 3 p.m.-8 p.m. 0.75% P 6 a.m.-11 a.m. 24.7a ±1.0 24.9a ±1.1 1.070a ± 0.0009 1.071a ±0.0009 1.50%P3p.m.-8p.m. 'P = Phosphorus. 2 3Values followed by different letters in the same column are significantly different (P < 0.05). S.E.M.
Serum calcium (mg. %) Egg specific gravity Treatment Exp. 3 Exp. 4 Exp. 3 Exp. 4 0,75% P16 a.m.-11 a.m. 29.7a2 ± 1.33 29.5a ±1.3 1.076a ± 0.0008 1.083a ± 0.0006 0.75% P 3 p.m.-8 p.m. 0.75% P 6 a.m.-11 a.m. 28.2a ±1.2 29.2a ±1.0 1.077a ± 0.0008 1.081a ± 0.0007 0.33%P3p.m.-8p.m. 'P = Phosphorus. 2 Values followed by different letters in the same column are significantly different (P < 0.05). 3 S.E.M. TABLE 3.—Serum calcium and egg specific gravity from hens fed various dietary treatments (Exps. 5 and 6) Serum calcium (mg. %) Egg specific gravity Treatment Exp. 5 Exp. 6 Exp. 5 Exp. 6 0.75%P 1 6a.m.-lla.m. 2 3 32.1a ± 0.8 32.1a ±2.1 1.075a ± 0.0006 1.079a ± 0.0008 0.75%P3p.m.-8p.m. 0.75%P6a.m.-ll a.m. 30.1a ±0.8 33.0a ±1.4 1,074a ± 0.0008 1.077a ± 0.0010 1.50% P 3 p.m.-8 p.m. 'P = Phosphorus. 2 3Values followed by different letters in the same column are significantly different (P < 0.05). S.E.M. (Taylor and Moore, 1956; Hurwitz and Bar, 1969; Roland et al., 1974). Aging hens producing many shell-less and thin shell eggs, however, do not appear to be obtaining maximum utilization of skeletal calcium since they have stronger bones and a greater bone ash than hens producing eggs with a better shell quality (Roland et al., 1975). The utilization of skeletal calcium for egg shell formation is apparently necessary and most important during the late hours of the night when total
calcium and percent calcium of the feed contents of the digestive system is at its lowest point (Roland era/., 1973). This release of calcium from the skeletal system is accompanied by a release of phosphorus which saturates the blood with phosphorus (Paul and Snetsinger, 1969). This may not be the best environment for a continued maximum stimulation of bone resorption necessary for optimal egg shell formation. It is postulated that the response of hens
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TABLE 2.—Serum calcium and egg specific gravity from hens fed various dietary treatments (Exps. 3 and 4)
C A : P RATIOS
in these experiments, although not always consistently different, in terms of specific gravity of eggs and serum calcium was due to the influence of the various Ca:P ratios on calcium absorption and/or bone resorption. ACKNOWLEDGEMENTS T h e s e investigations were supported in part b y a grant-in-aid from the Borden Chemical C o m p a n y (Smith-Douglas Division), Norfolk, Virginia.
Candlish, J. K., 1971. The formation of mineral and organic matrix of fowl cortical and medullary bone during shell calcification. Brit. Poultry Sci. 12: 119-127. Crowley, T. S., M. W. Pasvogel, A. R. Kemmerer, M. G. Vavich and A. A. Kurnick, 1961. Effects of soft phosphate and dicalcium phosphate on reproductive performance and egg quality. Poultry Sci. 40: 74-80. Cuisinier-Gleizes, P., A. George, C. Giuliano and H. Mathieu, 1971. Calcified Tissue: Structural, Functional and Metabolic Aspects. Eds. J. Menczel and A. Harrell. Academic Press Inc., N.Y., N.Y. Duncan, D. B., 1955. Multiple range and multiple F tests. Biometrics, 11: 1-42. Evans, R. J., J. S. Carver and A. W. Brant, 1944. The influence of dietary factors on eggshell quality. I. Phosphorus. Poultry Sci. 23: 9-15. Gillis, M. B., L. C. Norris and G. F. Heuser, 1953. Phosphorus metabolism and requirements of hens. Poultry Sci. 32: 977-984. Holcombe, D, J., D. A. Roland, Sr. and R. H. Harms, 1974. The ability of hens to adjust calcium or phosphorus intake when given a choice of diets containing different levels of either calcium or phosphorus. Poultry Sci. 53: 1936. Holcombe, D. J., D. A. Roland, Sr. and R. H. Harms, 1975. The ability of hens to adjust calcium intake when given diets containing two levels of calcium. Poultry Sci. 54: In press. Hurwitz, S., and A. Bar, 1969. Calcium reserves in bones of laying hens: Their presence and utilization. Poultry Sci. 48: 1391-1396. Morrissey,R. L.,andR.H. Wasserman, 1971. Calcium
absorption and calcium-binding protein in chicks on differing calcium and phosphorus intakes. Amer. J. Physiol. 220: 1509-1515. Paul, H. S., and D. C. Snetsinger, 1969. Dietary calcium and phosphorus and variations in plasma alkaline phosphatase activity in relationship to physical characteristics of eggshells. Poultry Sci. 48: 241-251. Raisz.L. C.,andl. Nieman, 1969. Effect of phosphate, calcium and magnesium on bone resorption and hormonal response in tissue culture. Endocrinology, 85: 446-452. Roland, D. A., Sr., D. J. Holcombe and R. H. Harms, 1975. Unpublished data. Fla. Agr. Exp. Sta. Roland, D. A., Sr., D. R. Sloan and R. H. Harms, 1973. Calcium metabolism in the laying hen. 4. The calcium status of the hen at night. Poultry Sci. 52: 351-354. Roland, D. A., Sr., D. R. Sloan, H. R. Wilson and R. H. Harms, 1974. Relationship of calcium to reproductive abnormalities in the laying hen (Gallus Domesticus). J. Nutr. 104: 1079-1085. Singsen, E. P., A. H. Spandorf, L. D. Matterson, J. A. Seratin and J. J. Tlustohowicz, 1962. Phosphorus in the nutrition of the adult hen. 1. Minimum phosphorus requirements. Poultry Sci. 41: 14011414. Snedecor, G. W., 1956. Statistical Methods, 5th edition. Iowa State University Press, Ames, Iowa. Taylor, T. G., 1970. The role of the skeleton in eggshell formation. Annales de Biologie Animale Biochimie Biophysique, 10: 83-90. Taylor, T. G., and J. H. Moore, 1956. The effect of calcium depletion on the chemical composition of bone minerals in laying hens. Brit. J. Nutr. 10: 250-263. Taylor, T. G., and J. H. Moore, 1958. The effect of high and low levels of dietary inorganic phosphate on the pre-laying storage of calcium and phosphorus and on the composition of the medullary and cortical bone in pullets. Brit. J. Nutr. 12: 35-42. Tyler, C , 1940a. Studies of calcium and phosphorus metabolism in relation to the chemical structure of bone. 1. Experiments with laying birds. Biochem. J. 34: 202-212. Tyler, C , 1940b. Studies of calcium and phosphorus metabolism in relation to the chemical structure of bone. 2. Experiments with molting birds. Biochem. J. 34: 1427-1430. Walter, E. D., and J. R. Aitken, 1962. Phosphorus requirements of laying hens confined to cages. Poultry Sci. 41: 386-390.
AUGUST 15-18, 1976. SIXTY-EIGHTH ANNUAL MEETING OF THE AMERICAN SOCIETY OF ANIMAL SCIENCE, TEXAS A AND M UNIVERSITY, COLLEGE STATION, TEXAS
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REFERENCES
641
Sharby, T. F., and J. M. Bell, 1974. In: Research on Rapeseed Seed, Oil and Meal. 3rd Progress Rep. Res. Committee, Rapeseed Assoc, Canada. Steel, R. G. D., and J. H. Torrie, 1960. Principles and Procedures of Statistics. McGraw-Hill Book Company Inc., Toronto. Stutz, M. W., J. E. Savage and B. L. O'Dell, 1971. Relation of dietary cations to arginine-lysine antagonism and free amino acid patterns in chicks. J. Nutr. 101: 377-394. Tao, R., R. J. Belzile and G. J. Brisson, 1971. Amino acid digestibility of rapeseed meal fed to chickens: effect of fat and lysine supplementation. Can. J. Anim. Sci. 51: 705-709. Vapar.Z., and D. R. Clandinin, 1972. Effect of tannins in rapeseed meal on its nutritional value for chicks. Poultry Sci., 51: 222-228. Williams, R. T., 1959. Cited in: Nesheim, M. C , and Garlich, J. D. 1963. Studies on ornithine synthesis in relation to benzoic acid excretion in the domestic fowl. J. Nutr. 79: 311-317.
The Influence of Feeding Diets Containing Different Calcium-Phosphorus Ratios on the Laying Hen1 D . A. ROLAND, S R . AND R. H . HARMS
Department ofPoultry Science, Florida Agricultural Experiment Station, Gainesville, Florida 32611 (Received for publication June 25, 1975)
ABSTRACT Six experiments were conducted to determine the influence of feeding diets containing different calcium-phosphorus (Ca:P) ratios at different times on the laying hen. Feeding a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 1.50% P from 3 p.m. to 8 p.m. decreased serum calcium in three of four experiments and decreased specific gravity of eggs in four of four experiments, when compared to that of the birds fed the control diet which contained 0.75% P during both time periods. Specific gravity of eggs from hens fed a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 0.33% P from 3 p.m. to 8 p.m. was increased in three of four experiments. Although the differences are not significant, with one exception, the trend in the experiments indicate that the level of P in the diet and time of intake can influence shell quality and serum calcium. The dietary treatments had no significant influence on feed consumption, egg production or egg weight during any of these experiments. It was postulated that the adverse or beneficial effect on specific gravity of eggs and serum calcium as a result of feeding diets containing various Ca:P ratios was due to the influence on calcium absorption and/or bone resorption. POULTRY SCIENCE 55: 637-641, 1976
INTRODUCTION
T
H E calcium-phosphorus (Ca:P) ratio is one of the best known concepts in poultry nutrition. Although it is extremely important in chick diets, it is generally believed that 1. Florida Agr. Exp. Sta. Journal Series No. 5972.
the C a : P ratio is not very important in laying h e n diets. M a n y studies h a v e been reported in which varying the p h o s p h o r u s c o n t e n t of the diet from 0.3 to 0.7% had no significant influence on egg shell quality (Crowley et al., 1961; E v a n s et al., 1944; Gillis et al., 1953; Singsen et al, 1962; Walter a n d Aitken,
Downloaded from http://ps.oxfordjournals.org/ at Oakland University on May 31, 2015
97-120 Pergomon Press, New York. Jones, J. D., 1964. Lysine-arginine antagonism in the chick. J. Nutr. 84: 313-321. Kadirvel, R., G. V. N. Rayudu and P. Vohra, 1969. Excretion of metabolites of tannic acid by chickens with and without ceca. Poultry Sci. 48: 1511-1513. Leslie, A. J., and J. D. Summers, 1975. Amino acid balance of rapeseed meal. Poultry Sci. 54: 532-537. Nesheim, M. C , and J. D. Garlich, 1963. Studies on ornithine synthesis in relation to benzoic acid excretion in the domestic fowl. J. Nutr. 79: 311. O'Dell, B. L., and J. E. Savage, 1966. Arginine-lysine antagonism in the chick and its relationship of dietary cations. J. Nutr. 90: 364-370. Olomu, J. M , A. R. Robblee and D. R. Clandinin, 1974. Effects of processing and amino acid supplementation on the nutritive value of rapeseed for broilers. Poultry Sci. 53: 175-184. Rayudu, G. V. N., R. Kadirvel, P. Vohra and F. H. Kratzer, 1970. Toxicity of tannic acid and its metabolites for chickens. Poultry Sci. 49: 957-960.
637
638
D. A. ROLAND, SR. AND R. H. HARMS
In the following studies, diets containing different Ca:P ratios were fed to hens at different periods during the day in an effort to create the best environment for maximum mobilization, utilization and repletion of skeletal calcium. EXPERIMENTAL PROCEDURE Experiment 1. One-hundred-and-twenty Babcock B-300 hens (hatched March 10, 1972)
were divided into three groups of 40 birds each on August 28, 1973 and fed one of the following treatments: The hens in group 1 were fed a control diet containing 0.75% phosphorus (P) from 6 a.m. to 11 a.m. and from 3 p.m. to 8 p.m. Hens in group 2 were fed a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 0.33% P from 3 p.m. to 8 p.m. (low phosphorus diet). Hens in group 3 were fed a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 1.50% P from 3 p.m. to 8 p.m. (high phosphorus diet). All diets contained 3.00% calcium and were iso-nitrogenous and iso-caloric. Hens were not given access to feed from 11 a.m. to 3 p.m. or from 8 p.m. to 6 a.m. Feed consumed during each feeding period was determined daily throughout the seven-day experiment. Specific gravity and egg weight were determined on all eggs. Blood was taken by anterior heart puncture from one-half of all birds at 6 a.m. on the eighth day and serum calcium determined. Experiment 2. The procedure of Experiment 2 was identical to Experiment 1 except that Babcock B-300 hens were hatched on May 22, 1973 and the experiment was initiated on October 29, 1974 and lasted for 10 days. Blood was taken at 6 a.m. on the 11th day. Experiment 3. One-hundred Babcock B-300 hens (hatched May 22, 1972) producing eggs with low specific gravity were divided into two groups. The hens were fed either a control diet containing 0.75% P from 6 a.m. to 11 a.m. and from 3 p.m. to 8 p.m. or a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 0.33% P from 3 p.m. to 8 p.m. Hens were not given access to feed from 11 a.m. to 3 p.m. or from 8 p.m. to 6 a.m. All diets contained 3.00% calcium and were iso-nitrogenous and iso-caloric. Feed consumed during each feeding period was determined daily throughout the 10-day experiment. Specific gravity and egg weight
Downloaded from http://ps.oxfordjournals.org/ at Oakland University on May 31, 2015
1962). More recent studies have indicated that phosphorus may have a more important role in calcium metabolism in the laying hen than previously believed. Holcombe et al. (1974) reported that the hen's tolerance of phosphorus was influenced by the calcium level of the diet and that extremely high levels of phosphorus had an adverse effect on shell quality. They also found that hens have the ability to adjust phosphorus intake when offered a choice of diets containing different levels of phosphorus. This ability of hens to regulate phosphorus intake was just as distinct as their ability to regulate calcium intake (Holcombe et al., 1975). Several recent studies have also indicated that phosphorus has a pronounced effect on calcium absorption and bone resorption (Raisz and Nieman, 1969; Morrissey and Wasserman, 1971). Cuisinier-Gleizes et al. (1971) reported that low phosphorus diets resulted in an immediate increase in bone resorption. The utilization of skeletal calcium for egg shell formation is apparently necessary and most important during the late hours of the night when total calcium and percent calcium of the feed contents of the digestive system is at its lowest (Roland et al., 1973). This release of calcium from the skeletal system into the blood is accompanied by a release of phosphorus. The increased amount of recycled phosphorus coupled with dietary phosphorus in the blood at this time may not be the best environment for maximum stimulation of bone resorption which is necessary for egg shell formation.
C A : P RATIOS
were determined on all eggs. Blood was taken by anterior heart puncture from one-half of all birds at 6 a.m. on the 11th day. Experiment 4. The procedure of Experiment 4 was similar to Experiment 3 except that 50 hens producing eggs with a high specific gravity were used.
Experiment 6. The procedure of Experiment 6 was identical to Experiment 5 except that 50 hens producing eggs with high specific gravity were used. The hens were fed either a diet containing 0.75% P from 6 a.m. to 11 a.m. and from 3 p.m. to 8 a.m. or a diet containing 0.75% P from 6 a.m. to 11 a.m. and a diet containing 1.50% P from 3 p.m. to 8 p.m. In all experiments serum calcium was determined using an atomic absorption spectrophotometer. All data were statistically analyzed by analysis of variance (Snedecor, 1956), and significant differences were determined by the multiple range test of Duncan (1955). RESULTS AND DISCUSSION In three of four experiments a decrease in serum calcium occurred in hens fed the diet containing 1.50% P from 3 p.m. to 8 p.m. (Tables 1 and 3). The decrease in serum calcium approached significance at the 0.05 level in Experiments 1, 2 and 5. Serum calcium of hens fed the diet containing 0.33% P from 3 p.m. to 8 p.m. in Experiments 1-4 was numerically reduced (Tables 1 and 2).
Specific gravity of eggs from hens fed the diet containing 1.50% P from 3 p.m. to 8 p.m. was decreased in four of four experiments (Tables 1 and 3). Specific gravity of eggs from hens fed the diet containing 0.33% P from 3 p.m. to 8 p.m. was increased in three of four experiments (Tables 1 and 2). Dietary treatment had no significant influence on feed consumption, egg production or egg weight during the experiments. The hens consumed an average of 53% more feed from 3 p.m. to 8 p.m. than from 6 a.m. to 11 a.m. Feeding hens a diet containing a 2:1 Ca:P ratio from 3 p.m. to 8 p.m. appeared to have an adverse effect on serum calcium and specific gravity of eggs. However, feeding hens a diet containing a 9:1 Ca:P ratio from 3 p.m. to 8 p.m. had a beneficial effect on specific gravity of eggs in three of four experiments. In Experiment 2, specific gravity of eggs from hens fed the 9:1 Ca:P ratio during the evening was significantly greater than that from hens fed the diet containing a 2:1 Ca:P ratio during the same period. The results of these studies suggest that the Ca:P ratio of the diet fed to hens can influence serum calcium and specific gravity of eggs. It is believed that the response of hens in these experiments to various Ca:P ratios was due to the influence of phosphorus on calcium metabolism. Several studies have indicated that phosphorus has a pronounced effect on calcium absorption and bone resorption (Raisz and Nieman, 1969; Morrissey and Wasserman, 1971) Cuisinier-Gleizes et al. (1971) reported that low phosphorus diets resulted in an immediate increase in bone resorption. Several workers have also shown that calcium in the skeletal system is important in the process of egg shell formation (Tyler, 1940a, b; Taylor and Moore, 1958; Taylor, 1970; Candlish, 1971). Large quantities of calcium were mobilized from the skeletal system and utilized when calcium was not available from the intestinal contents
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Experiment 5. The procedure of Experiment 5 was similar to Experiment 3 except that Experiment 5 was initiated on April 15, 1974. The hens producing eggs with poor egg shell quality were fed either a diet containing 0.75% P from 6 a.m. to 11 a.m. and from 3 p.m. to 8 p.m. or a diet containing 0.33% P from 3 p.m. to 8 p.m.
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D. A. ROLAND, SR. AND R. H. HARMS
TABLE 1.—Serum calcium and egg specific gravity from hens fed various dietary treatments (Exps. 1 and 2) Serum calcium (mg. %) Egg specific gravity Treatment Exp. 1 Exp. 2 Exp. Exp. 2 0.75%P l 6a.m.-lla.m. 2 3 26.9a ± 1.2 27.0a ± 1.2 1.071a ± 0.0011 1.073ab ± 0.0010 0.75% P 3 p.m.-8 p.m. 0.75% P 6 a.m.-11 a.m. 26.8a ±1.0 26.6a ± 0.9 1.072a ± 0.0009 1.074b ±0.0009 0.33% P 3 p.m.-8 p.m. 0.75% P 6 a.m.-11 a.m. 24.7a ±1.0 24.9a ±1.1 1.070a ± 0.0009 1.071a ±0.0009 1.50%P3p.m.-8p.m. 'P = Phosphorus. 2 3Values followed by different letters in the same column are significantly different (P < 0.05). S.E.M.
Serum calcium (mg. %) Egg specific gravity Treatment Exp. 3 Exp. 4 Exp. 3 Exp. 4 0,75% P16 a.m.-11 a.m. 29.7a2 ± 1.33 29.5a ±1.3 1.076a ± 0.0008 1.083a ± 0.0006 0.75% P 3 p.m.-8 p.m. 0.75% P 6 a.m.-11 a.m. 28.2a ±1.2 29.2a ±1.0 1.077a ± 0.0008 1.081a ± 0.0007 0.33%P3p.m.-8p.m. 'P = Phosphorus. 2 Values followed by different letters in the same column are significantly different (P < 0.05). 3 S.E.M. TABLE 3.—Serum calcium and egg specific gravity from hens fed various dietary treatments (Exps. 5 and 6) Serum calcium (mg. %) Egg specific gravity Treatment Exp. 5 Exp. 6 Exp. 5 Exp. 6 0.75%P 1 6a.m.-lla.m. 2 3 32.1a ± 0.8 32.1a ±2.1 1.075a ± 0.0006 1.079a ± 0.0008 0.75%P3p.m.-8p.m. 0.75%P6a.m.-ll a.m. 30.1a ±0.8 33.0a ±1.4 1,074a ± 0.0008 1.077a ± 0.0010 1.50% P 3 p.m.-8 p.m. 'P = Phosphorus. 2 3Values followed by different letters in the same column are significantly different (P < 0.05). S.E.M. (Taylor and Moore, 1956; Hurwitz and Bar, 1969; Roland et al., 1974). Aging hens producing many shell-less and thin shell eggs, however, do not appear to be obtaining maximum utilization of skeletal calcium since they have stronger bones and a greater bone ash than hens producing eggs with a better shell quality (Roland et al., 1975). The utilization of skeletal calcium for egg shell formation is apparently necessary and most important during the late hours of the night when total
calcium and percent calcium of the feed contents of the digestive system is at its lowest point (Roland era/., 1973). This release of calcium from the skeletal system is accompanied by a release of phosphorus which saturates the blood with phosphorus (Paul and Snetsinger, 1969). This may not be the best environment for a continued maximum stimulation of bone resorption necessary for optimal egg shell formation. It is postulated that the response of hens
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TABLE 2.—Serum calcium and egg specific gravity from hens fed various dietary treatments (Exps. 3 and 4)
C A : P RATIOS
in these experiments, although not always consistently different, in terms of specific gravity of eggs and serum calcium was due to the influence of the various Ca:P ratios on calcium absorption and/or bone resorption. ACKNOWLEDGEMENTS T h e s e investigations were supported in part b y a grant-in-aid from the Borden Chemical C o m p a n y (Smith-Douglas Division), Norfolk, Virginia.
Candlish, J. K., 1971. The formation of mineral and organic matrix of fowl cortical and medullary bone during shell calcification. Brit. Poultry Sci. 12: 119-127. Crowley, T. S., M. W. Pasvogel, A. R. Kemmerer, M. G. Vavich and A. A. Kurnick, 1961. Effects of soft phosphate and dicalcium phosphate on reproductive performance and egg quality. Poultry Sci. 40: 74-80. Cuisinier-Gleizes, P., A. George, C. Giuliano and H. Mathieu, 1971. Calcified Tissue: Structural, Functional and Metabolic Aspects. Eds. J. Menczel and A. Harrell. Academic Press Inc., N.Y., N.Y. Duncan, D. B., 1955. Multiple range and multiple F tests. Biometrics, 11: 1-42. Evans, R. J., J. S. Carver and A. W. Brant, 1944. The influence of dietary factors on eggshell quality. I. Phosphorus. Poultry Sci. 23: 9-15. Gillis, M. B., L. C. Norris and G. F. Heuser, 1953. Phosphorus metabolism and requirements of hens. Poultry Sci. 32: 977-984. Holcombe, D, J., D. A. Roland, Sr. and R. H. Harms, 1974. The ability of hens to adjust calcium or phosphorus intake when given a choice of diets containing different levels of either calcium or phosphorus. Poultry Sci. 53: 1936. Holcombe, D. J., D. A. Roland, Sr. and R. H. Harms, 1975. The ability of hens to adjust calcium intake when given diets containing two levels of calcium. Poultry Sci. 54: In press. Hurwitz, S., and A. Bar, 1969. Calcium reserves in bones of laying hens: Their presence and utilization. Poultry Sci. 48: 1391-1396. Morrissey,R. L.,andR.H. Wasserman, 1971. Calcium
absorption and calcium-binding protein in chicks on differing calcium and phosphorus intakes. Amer. J. Physiol. 220: 1509-1515. Paul, H. S., and D. C. Snetsinger, 1969. Dietary calcium and phosphorus and variations in plasma alkaline phosphatase activity in relationship to physical characteristics of eggshells. Poultry Sci. 48: 241-251. Raisz.L. C.,andl. Nieman, 1969. Effect of phosphate, calcium and magnesium on bone resorption and hormonal response in tissue culture. Endocrinology, 85: 446-452. Roland, D. A., Sr., D. J. Holcombe and R. H. Harms, 1975. Unpublished data. Fla. Agr. Exp. Sta. Roland, D. A., Sr., D. R. Sloan and R. H. Harms, 1973. Calcium metabolism in the laying hen. 4. The calcium status of the hen at night. Poultry Sci. 52: 351-354. Roland, D. A., Sr., D. R. Sloan, H. R. Wilson and R. H. Harms, 1974. Relationship of calcium to reproductive abnormalities in the laying hen (Gallus Domesticus). J. Nutr. 104: 1079-1085. Singsen, E. P., A. H. Spandorf, L. D. Matterson, J. A. Seratin and J. J. Tlustohowicz, 1962. Phosphorus in the nutrition of the adult hen. 1. Minimum phosphorus requirements. Poultry Sci. 41: 14011414. Snedecor, G. W., 1956. Statistical Methods, 5th edition. Iowa State University Press, Ames, Iowa. Taylor, T. G., 1970. The role of the skeleton in eggshell formation. Annales de Biologie Animale Biochimie Biophysique, 10: 83-90. Taylor, T. G., and J. H. Moore, 1956. The effect of calcium depletion on the chemical composition of bone minerals in laying hens. Brit. J. Nutr. 10: 250-263. Taylor, T. G., and J. H. Moore, 1958. The effect of high and low levels of dietary inorganic phosphate on the pre-laying storage of calcium and phosphorus and on the composition of the medullary and cortical bone in pullets. Brit. J. Nutr. 12: 35-42. Tyler, C , 1940a. Studies of calcium and phosphorus metabolism in relation to the chemical structure of bone. 1. Experiments with laying birds. Biochem. J. 34: 202-212. Tyler, C , 1940b. Studies of calcium and phosphorus metabolism in relation to the chemical structure of bone. 2. Experiments with molting birds. Biochem. J. 34: 1427-1430. Walter, E. D., and J. R. Aitken, 1962. Phosphorus requirements of laying hens confined to cages. Poultry Sci. 41: 386-390.
AUGUST 15-18, 1976. SIXTY-EIGHTH ANNUAL MEETING OF THE AMERICAN SOCIETY OF ANIMAL SCIENCE, TEXAS A AND M UNIVERSITY, COLLEGE STATION, TEXAS
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REFERENCES
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