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Gross Abnormalities in Chicks Fed Amino Acid Deficient Diets1
Gross Abnormalities in Chicks Fed Amino Acid Deficient Diets1 J. O. ANDERSON AND R. E. WARNICK Animal Science Department, Utah State University, Logan, Utah 84321 (Received for publication November 8, 1966)
E
XPERIMENTS are being conducted at this station to determine by chick tests which amino acids are most limiting in the protein of certain feedstuffs used in commercial poultry rations. In our recent tests, a commercial feedstuff was used to provide 14 percent protein in the semipurified rations fed. The feedstuff was supplemented with an amino acid mixture that gave an 18 percent protein basic ration with a balanced mixture of the essential amino acids. The amino acid mixture provided a greater fraction of the chick's requirement for a given amino acid when the feedstuff being studied was relatively more deficient in that amino acid. Each essential amino acid in the basic ration's amino acid mixture was removed in turn to produce the other rations fed. Chick growth rate was decreased most by removal of the amino acids most deficient in the feedstuff being studied. Abnormal feather development was noted when certain amino acids were removed. Other experiments were then conducted to determine what deficiency symptoms would be produced when the degree of deficiency, as judged by growth rate, was approximately the same for each of the essential amino acids. This paper describes the results obtained in these later experiments. Several researchers have described abnormal feathering that was corrected by the addition of more arginine to the diet. "Approved as Utah Agricultural Experiment Station Journal Paper No. 616.
Hegsted et al. (1941) reported that the feathers of chicks fed arginine-deficient diets were more brittle and easily broken, they had a thin and ragged appearance, and the barbs were less well developed. A paralysis noted among the deficient birds was characterized by a high-stepping stilted gait with hocks thrust forward and toes extended. The muscles of these birds were poorly developed. Sanders et al. (1950) fed chicks a ration based on milo gluten meal and lysine. The chick's primaries and secondaries had a spoon-like appearance caused by the retention of an abnormally long sheath that covered the proximal onethird of the feather shaft. The barbs and barbules were limp and degenerate on the part of the shaft next to that covered by the sheath. The integrity of the vane was maintained by normal barbs only at the tip of the feather. The rachis of each primary and secondary was bent abnormally. These abnormal conditions were not noted when arginine was added to the diet. Several other groups have described somewhat similar conditions in chicks fed arginine deficient diets. Fritz et al. (1946) reported that Bronze turkey poults fed rations high in certain vegetable proteins failed to develop normal feather pigmentation in the flight feathers. This syndrome was prevented by adding lysine or proteins rich in lysine to the diet. These investigators indicated that some of the poults also developed a curled feather condition. Several other reports on this condition in turkeys have appeared since 1946. Klain et al. (1957) found that chicks of
856
AMINO ACID DEFICIENCY
four different breeds developed a similar achromatosis of the feathers when they were fed a lysine deficient diet. This symptom was not prevented by the addition of folic acid, iodinated casein, or tyrosine to the diet, although tyrosine addition appeared to reduce the severity of the achromatosis. Brief notes have appeared in the literature on feather abnormalities found after feeding rations deficient in other essential amino acids. Klain et al. (1960) reported that chicks fed a ration with 0.66 percent L-leucine developed twisted and ruffled feathers. Wilkening et al. (1947) noted ragged primaries among chicks fed a tryptophan deficient diet. Grau (1945) reported that chicks fed rations deficient in isoleucine, leucine, or phenylalanine developed a folded tongue. Ousterhout (1960) fed chicks rations lacking each of the essential amino acids. Folded tongues were noted when chicks were fed rations lacking any one of seven of the essential amino acids. PROCEDURE
The composition of the two basic rations fed in these experiments is given in Table 1. Ration 1 was based on a mixture of amino acids; the essential amino acids were present in the mixture at the levels found by Dobson et al. (1964) to be well balanced (see Table 2 of this paper). The racemic form of four amino acids was used in most rations. However, rations deficient in valine, isoleucine, or threonine contained only the natural isomer of the deficient amino acid. The deficient diets were produced by removing approximately 46 percent of the basic ration's level of each essential amino acid with only one amino acid being reduced in each deficient ration. Ration 2 was based on a mixture of herring meal and amino acids. The calculated level of each essential amino acid was ap-
857
SYMPTOMS
TABLE 1.—Composition of the basic rations Ingredients Herring meal, 71.9% prot. L-arginine L-arginine HC1 L-histidineHCl-H 2 0 L-lysine HC1 L-leucine DL-isoleucine DL-valine L-phenylalanine L-tryosine L-tryptophan DL-methionine L-cystine DL-threonine Glycine L-proline L-asparagine Sodium glutamate Potassium glutamate L-glutamic acid Vitamin mix1 Dicalcium phosphate 2 Calcium carbonate Disodium phosphate Magnesium sulfate Magnesium trisilicate Aluminum hydroxide Trace mineral mix3 Salt Cottonseed oil Cellulose4 Starch to Glucose monohydrate 6 to
Ration 1
Ration 2
gms./kg. 139
—
3 11.9 5.8 14.35 13 16 19 6.7 6.6 2 (DL) 3.9 3.4 15.6 20 6 7.5 11.55 17.65 40.6 100 28.5 10 7 3 6 3 0.6
—
40 20 1,000
—
—
6.3 2.4 4 5.1 6.8 7 2.7 3.3 1.6 1.2 1.9 6.5 5 1
—
12.2 9.2 35.5 80 20 3
—
2.1
— —
0.6 1.5 15 25
—
1,000
1
Composition given by Dobson et al. (1964). A purified dicalcium phosphate was used in ration 1 and a feed grade was used in ration 2 (18.5% 2
P
)'3
.
.
.
This mix provided the following in gms./kg. of ration: FeC 6 H 5 07, 0.2; MnS0 4 -H 2 0, 0.25; K I 0 3 ) 0.01; ZnCOs, 0.13; and CuS0 4 , 0.01. 4 Alphacel, Nutritional Biochemicals Corporation, Cleveland, Ohio. 6 Cerelose, Corn Products Company, New York, New York.
proximately the same as in ration 1. Deficiencies were produced by removing amino acids from the amino acid mixture and varied in degree according to level of each amino acid in herring meal. More glutamic acid was added to keep the nitrogen level constant when the level of an essential amino acid was reduced in ration 1 or 2. Two white and two colored chicks were fed each ration in each experiment. The white chicks were from a White Vantress X
858
J. 0 . ANDERSON AND R. E. WARNICK TABLE 2.—Mean weight gains obtained when the level of each amino acid was reduced Ration 1
Amino acid reduced
None Arginine Histidine Lysine Leucine Isoleucine Valine Phenylalanine & tyrosine Tryptophan Methionine & cystine Threonine
Ration 2
Reduced level1
Gain, gms./day
No. birds
—
11.0 4.5 5.5 5.5 4.5 4.0 4.0 4.5
16 4 12 8 8 8 8 8
ro.20
5.0 5.0
8 4
\0.18 0.43
5.0
12
0.70 0.22 0.64 0.70 0.45 0.50 JO. 37 \0.36 0.11
Reduced level2
Gain, gms./day
No. birds
0.78
11.0 5.0
12 4
—
—
—
4.5 4.0
8 4
—
—
0.82 0.72 0.42 0.79 0.36 0.28 0.10 0.27 0.10
—
8.0 5.0 4.0 4.0 4.0
4 4 8 4 16
1
Essential amino acid levels in ration 1 were as follows (in percent): arginine, 1.28; histidine, 0.43; lysine, 1.15; leucine, 1.30; isoleucine, 0.80; valine, 0.95; phenylalanine, 0.67; tyrosine, 0.66; tryptophan, 0.20; methionine, 0.38; cystine, 0.35; and threonine, 0.78. The D-isomer of isoleucine, valine, and threonine was considered to be inactive. 2 Based on values given by March el al. (1962).
White Plymouth mating. The colored chicks were produced by mating White Plymouth Rock females with colored males produced from a Red Vantress X White Plymouth Rock mating. A variety of colors and color patterns was found among the colored chicks. The chicks were not sexed. The chicks were reared in electricallyheated batteries with wire floors. Feed and water were supplied ad libitum. Chicks were fed a practical type broiler ration for one day after being taken from the incubator and then distributed among experimental groups on the basis of body weight and color. Experimental rations were fed for approximately two weeks and the birds were then returned to the practical ration. Pictures were taken about one week later to allow some of the modified feathers to develop to a point where the modifications were visible. RESULTS AND DISCUSSION
The mean weight gains are summarized in Table 2. Weight gains were reduced from 11.0 to from 4.0 to 5.5 grams per day by modifying ration 1 and 2. Herring meal
is an excellent source of lysine, so the mean weight gain was not reduced as much when this amino acid was removed from the amino acid mixture used with ration 2. Large numbers of chicks were not fed each ration in these tests. Most of the changes in feather structure noted were evident after the first experiment. Lysine, threonine, and methionine and cystine deficient rations were fed in our studies on peanut meal (Anderson and Warnick, 1965) and cottonseed meal (Anderson and Warnick, 1966), so it did not seem justified to feed additional chicks deficient rations. The observations made in this report are based primarily on the experiments conducted with rations 1 and 2. Some statements, however, necessarily also draw upon observations made during the above listed experiments. Abnormal feather development was noted when arginine, isoleucine, leucine, valine, or phenylalanine and tyrosine were deficient in the diet. Chicks fed the tryptophan deficient diet showed some ragged feathering but not to the degree found among chicks fed the diets with the five
AMINO ACID DEFICIENCY SYMPTOMS
859
FIG. 1. Chick on the right was fed ration 1, while the chick on the left was fed ration 1 with the valine level reduced to 0.5 percent.
deficiencies listed first. When the ration was deficient in histidine, lysine, threonine, or methionine and cystine, growth rate and feather development were slower than with the basic ration, but the chicks appeared normal otherwise. Even the achromatosis usually associated with lysine deficiency did not appear when part of the lysine was removed from rations 1 and 2. This lack of pigment had been noted several times when other lysine deficient rations were fed in our laboratory. Abnormal feather development had not come to our attention in our earlier experiments in which diets deficient in histidine, threonine, or methionine and cystine were fed. However, abnormal feathering may develop if rations containing more serious deficiencies of these amino acids are fed. The abnormal feather conditions noted among chicks fed rations deficient in arginine, valine, leucine, isoleucine, tryptophan, or phenylalanine and tyrosine were somewhat similar. If there was a difference between any two deficiencies it was only in the degree to which the abnormalities de-
veloped. Similar symptoms were produced when either ration 1 or 2 was made deficient in one of the amino acids. The abnormalities noted included the ragged appearance reported by Hegsted et al. (1941) and the spoon-shaped and curled feathers noted by Sanders et al. (1950). Often the rachis of each secondary was found to curl toward the body when the wing was held in the open position. The small tracts of feathers on the shoulders curled upwards; these changes were noted within a few days after the chicks were started on the deficient rations. Many feathers lacked their characteristic smooth surface. When these feathers were examined under the microscope it was found that the barbules on adjacent barbs did not form the normal interlocking structure, and the barbules did not spread as much as in normal feathers. Hooklets were usually present although, in some cases, the barbules appeared to have been broken off with a loss of the hooklets. These conditions gave the feathers a ragged and sometimes thin appearance.
860
J. O. ANDERSON AND R. E. WARNICK
FIC. 2. Chick fed ration 2 with the arginine removed from the amino acid mixture.
The chick on the right in Figure 1 was fed ration 1 during the experimental period. Chicks fed this ration were not as smoothly feathered as chicks fed a well balanced broiler ration, but their feathering was smoother than that of chicks fed any of the six deficient rations listed above. The
FIG. 3. Chick fed ration 1 with the leucine level reduced to 0.7 percent.
chick on the left in Figure 1 was fed ration 1 with the valine level reduced from 0.95 to 0.5 percent. This chick was typical of the valine deficient birds. The feathering was ragged and the secondaries showed a marked curl. When the wing was held in the normal position the secondaries extended above the bird's backline. The chick in Figure 2 was fed ration 2 with the arginine removed from the amino acid mixture. Fewer than 25 percent of the birds fed this ration were affected to this extent. The feather condition was similar to that described by Hegsted et al. (1941) and Sanders et al. (1950). Some of the arginine deficient birds were unsteady while they were standing. Weak birds were noted in groups of chicks fed rations deficient in other amino acids, but the unsteadiness while standing and the unusual gait seemed to be more common or even peculiar to the arginine deficient chicks. Some birds fed this ration appeared normal except for the smaller body size. The chick in Figure 3 was fed ration 1 and the leucine level reduced from 1.3 to 0.7 percent. The feathering of this chick was affected to a greater degree than that of the average bird fed the leucine deficient rations. The leucine deficient chicks showed
FIG. 4. Chick fed ration 1 with the isoleucine level reduced to 0.45 percent.
AMINO ACID DEFICIENCY SYMPTOMS
more ragged, curled, and twisted feathers. Even the primary and secondary coverts and the breast feathers of the pictured chick were curled abnormally. The chick in Figure 4 was fed ration 1 wtih isoleucine level reduced from 0.8 to 0.4S percent. The feathering of birds fed this ration was less affected than that of the birds fed the valine, arginine, or leucine deficient diets. About one-third of these birds showed flight feathers with a ragged and thin appearance. Colored chicks fed the rations deficient in phenylalanine and tyrosine showed changes in the feather pigments (see Figure 5). The most obvious difference was noted with black feathered chicks. These chicks showed a gray bar across the flight feathers; the color of body feathers was also affected. Red birds showed some less obvious loss of pigments. Both colored and white chicks showed ragged feathering. SUMMARY
Chicks were fed rations deficient in each of the essential amino acids for two weeks. The deficient rations were produced by removing in turn approximately 46 percent of each essential amino acid from rations with a well balanced mixture of the essential amino acids. Reducing the essential amino acid levels reduced the mean growth
FIG. 5. Chick fed ration 2 without the phenylalanine and tyrosine of the amino acid mixture.
861
rate from 11 to approximately 4.S grams per day. Abnormal feather structures were evident when the ration was deficient in leucine, arginine, valine, isoleucine, tryptophan, or phenylalanine and tyrosine. Abnormalities included curled feathers, spoonshaped feathers, and feathers with a ragged and thin appearance. The feather abnormalities produced with each of the six listed deficiencies were somewhat similar, but these symptoms developed to different degrees. Reduced feather pigmentation was noted when a phenylalanine and typrosine deficient ration was fed to colored chicks. REFERENCES Anderson, J. O., and R. E. Warnick, 1965. Amino acid deficiencies in peanut meal and in corn and peanut meal rations. Poultry Sci. 44: 10661072. Anderson J. O., and R. E. Warnick, 1966. Sequence in which essential amino acids become limiting for growth of chicks fed rations containing cottonseed meal. Poultry Sci. 45: 8489. Dobson, D. C , J. O. Anderson and R. E. Warnick, 1964. A determination of the essential amino acid proportions needed to allow rapid growth in chicks. J. Nutrition, 82: 67-75. Fritz, J. C , J. H. Hooper, J. L. Halpin and H. P. Moore, 1946. Failure of feather pigmentation in bronze poults due to lysine deficiency. J. Nutrition, 3 1 : 387-396. Grau, C. R., 1945. Deformity of the tongue associated with amino acid deficiencies in the chick. Proc. Soc. Exp. Biol. Med. 59: 177-178. Hegsted, D. M., G. M. Briggs, C. A. Elvehjem and E. B. Hart, 1941. The role of arginine and glycine in chick nutrition. J. Biol. Chem. 140: 191-200. Klain, G. J., D. C. Hill, J. A. Gray and H. D. Branion, 1957. Achromatosis in the feathers of chicks fed lysine-deficient diets. J. Nutrition, 61 : 317-328. Klain, G. J., H. M. Scott and B. C. Johnson, 1960. The amino acid requirements of the growing chick fed a crystalline amino acid diet. Poultry Sci. 39 : 39-44. March, B. E., J. Biely and H. L. A. Tarr, 1962. Nutrient composition of British Columbia herring meal. Fisheries Research Board of Canada Circular No. 26.
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J. O. ANDERSON AND R. E.
Ousterhout, L. E., 1960. Survival time and biochemical changes in chicks fed diets lacking different essential amino acids. J. Nutrition, 70: 226-234. Sanders, B. G., S. O. Brown and J. R. Couch, 1950. A feathering syndrome in chicks after feeding optimal levels of lysine in the absence
WAENICK
of arginine. Proc. Soc. Exp. Biol. Med. 74: 114— 117. Wilkening, M. C , B. S. Schweigert, P. B. Pearson and R. M. Sherwood, 1947. Studies on the requirement of the chick for tryptophan. J. Nutrition, 34: 701-714.
The Utilization of Phytate Phosphorus by Poultry—A Review T. S. N E L S O N
Research and Development Division, International Minerals and Chemical Libertyville, Illinois 60048
Corporation,
(Received for publication November 8, 1966)
T
HE metabolism of phosphorus derived from plant tissues is one of the least understood and most debated subjects in the field of mineral nutrition. A small proportion of the total phosphorus in plants is inorganic phosphates located primarily in the vegetative tissues. Most of the phosphorus is in a variety of organic compounds found primarily in the seed, and phytate phosphorus is the predominant organic form. The amount of phytate phosphorus utilized by animals has economic importance because seeds are the major plant-source ingredients used in feeds. Many scientists currently use the rule that monogastric animals metabolize only one-third of the phosphorus in plant materials. This assumption is based on the report by the Committee on Animal Nutrition, National Academy of Sciences-National Research Council (NAS-NRC) (1960) that approximately 30% of the phosphorus in plant materials in non-phytate and can be considered to be utilized by animals. Many scientists using a variety of experimental procedures have studied the ability of different species of animals to utilize phytate phosphorus. However, they have failed to define to everyone's satisfaction
the extent of phytate phosphorus availability in strictly quantitative terms. Certain authors have reported it was utilized to a limited extent. Others have considered it was highly available to animals. Kastelic and Forbes (1961) and Taylor (1965) reviewed this subject. They stressed that there is still no general agreement on the extent to which different species of animals at various ages utilize phytate phosphorus. Studies of Phytate Phosphorus Utilization Heuser et al. (1943), McGinnis et al. (1944), Singsen et al. (1947), Gillis et al. (1949), and Sunde and Bird (1956) observed that natural phytate was a poor source of phosphorus for various species of poultry. Conversely, Sieburth et al. (1952) reported the phosphorus in finely ground whole wheat flour was almost completely available to chicks for growth but was less available than inorganic phosphate for bone deposition. Temperton et al. (1965a, b, c) concluded that pullet chicks less than four weeks old, growing pullets reared to 18 weeks of age and laying hens were able to achieve effective utilization of organic sources of phosphorus for growth and bone formation. Several investigators have fed various
E
XPERIMENTS are being conducted at this station to determine by chick tests which amino acids are most limiting in the protein of certain feedstuffs used in commercial poultry rations. In our recent tests, a commercial feedstuff was used to provide 14 percent protein in the semipurified rations fed. The feedstuff was supplemented with an amino acid mixture that gave an 18 percent protein basic ration with a balanced mixture of the essential amino acids. The amino acid mixture provided a greater fraction of the chick's requirement for a given amino acid when the feedstuff being studied was relatively more deficient in that amino acid. Each essential amino acid in the basic ration's amino acid mixture was removed in turn to produce the other rations fed. Chick growth rate was decreased most by removal of the amino acids most deficient in the feedstuff being studied. Abnormal feather development was noted when certain amino acids were removed. Other experiments were then conducted to determine what deficiency symptoms would be produced when the degree of deficiency, as judged by growth rate, was approximately the same for each of the essential amino acids. This paper describes the results obtained in these later experiments. Several researchers have described abnormal feathering that was corrected by the addition of more arginine to the diet. "Approved as Utah Agricultural Experiment Station Journal Paper No. 616.
Hegsted et al. (1941) reported that the feathers of chicks fed arginine-deficient diets were more brittle and easily broken, they had a thin and ragged appearance, and the barbs were less well developed. A paralysis noted among the deficient birds was characterized by a high-stepping stilted gait with hocks thrust forward and toes extended. The muscles of these birds were poorly developed. Sanders et al. (1950) fed chicks a ration based on milo gluten meal and lysine. The chick's primaries and secondaries had a spoon-like appearance caused by the retention of an abnormally long sheath that covered the proximal onethird of the feather shaft. The barbs and barbules were limp and degenerate on the part of the shaft next to that covered by the sheath. The integrity of the vane was maintained by normal barbs only at the tip of the feather. The rachis of each primary and secondary was bent abnormally. These abnormal conditions were not noted when arginine was added to the diet. Several other groups have described somewhat similar conditions in chicks fed arginine deficient diets. Fritz et al. (1946) reported that Bronze turkey poults fed rations high in certain vegetable proteins failed to develop normal feather pigmentation in the flight feathers. This syndrome was prevented by adding lysine or proteins rich in lysine to the diet. These investigators indicated that some of the poults also developed a curled feather condition. Several other reports on this condition in turkeys have appeared since 1946. Klain et al. (1957) found that chicks of
856
AMINO ACID DEFICIENCY
four different breeds developed a similar achromatosis of the feathers when they were fed a lysine deficient diet. This symptom was not prevented by the addition of folic acid, iodinated casein, or tyrosine to the diet, although tyrosine addition appeared to reduce the severity of the achromatosis. Brief notes have appeared in the literature on feather abnormalities found after feeding rations deficient in other essential amino acids. Klain et al. (1960) reported that chicks fed a ration with 0.66 percent L-leucine developed twisted and ruffled feathers. Wilkening et al. (1947) noted ragged primaries among chicks fed a tryptophan deficient diet. Grau (1945) reported that chicks fed rations deficient in isoleucine, leucine, or phenylalanine developed a folded tongue. Ousterhout (1960) fed chicks rations lacking each of the essential amino acids. Folded tongues were noted when chicks were fed rations lacking any one of seven of the essential amino acids. PROCEDURE
The composition of the two basic rations fed in these experiments is given in Table 1. Ration 1 was based on a mixture of amino acids; the essential amino acids were present in the mixture at the levels found by Dobson et al. (1964) to be well balanced (see Table 2 of this paper). The racemic form of four amino acids was used in most rations. However, rations deficient in valine, isoleucine, or threonine contained only the natural isomer of the deficient amino acid. The deficient diets were produced by removing approximately 46 percent of the basic ration's level of each essential amino acid with only one amino acid being reduced in each deficient ration. Ration 2 was based on a mixture of herring meal and amino acids. The calculated level of each essential amino acid was ap-
857
SYMPTOMS
TABLE 1.—Composition of the basic rations Ingredients Herring meal, 71.9% prot. L-arginine L-arginine HC1 L-histidineHCl-H 2 0 L-lysine HC1 L-leucine DL-isoleucine DL-valine L-phenylalanine L-tryosine L-tryptophan DL-methionine L-cystine DL-threonine Glycine L-proline L-asparagine Sodium glutamate Potassium glutamate L-glutamic acid Vitamin mix1 Dicalcium phosphate 2 Calcium carbonate Disodium phosphate Magnesium sulfate Magnesium trisilicate Aluminum hydroxide Trace mineral mix3 Salt Cottonseed oil Cellulose4 Starch to Glucose monohydrate 6 to
Ration 1
Ration 2
gms./kg. 139
—
3 11.9 5.8 14.35 13 16 19 6.7 6.6 2 (DL) 3.9 3.4 15.6 20 6 7.5 11.55 17.65 40.6 100 28.5 10 7 3 6 3 0.6
—
40 20 1,000
—
—
6.3 2.4 4 5.1 6.8 7 2.7 3.3 1.6 1.2 1.9 6.5 5 1
—
12.2 9.2 35.5 80 20 3
—
2.1
— —
0.6 1.5 15 25
—
1,000
1
Composition given by Dobson et al. (1964). A purified dicalcium phosphate was used in ration 1 and a feed grade was used in ration 2 (18.5% 2
P
)'3
.
.
.
This mix provided the following in gms./kg. of ration: FeC 6 H 5 07, 0.2; MnS0 4 -H 2 0, 0.25; K I 0 3 ) 0.01; ZnCOs, 0.13; and CuS0 4 , 0.01. 4 Alphacel, Nutritional Biochemicals Corporation, Cleveland, Ohio. 6 Cerelose, Corn Products Company, New York, New York.
proximately the same as in ration 1. Deficiencies were produced by removing amino acids from the amino acid mixture and varied in degree according to level of each amino acid in herring meal. More glutamic acid was added to keep the nitrogen level constant when the level of an essential amino acid was reduced in ration 1 or 2. Two white and two colored chicks were fed each ration in each experiment. The white chicks were from a White Vantress X
858
J. 0 . ANDERSON AND R. E. WARNICK TABLE 2.—Mean weight gains obtained when the level of each amino acid was reduced Ration 1
Amino acid reduced
None Arginine Histidine Lysine Leucine Isoleucine Valine Phenylalanine & tyrosine Tryptophan Methionine & cystine Threonine
Ration 2
Reduced level1
Gain, gms./day
No. birds
—
11.0 4.5 5.5 5.5 4.5 4.0 4.0 4.5
16 4 12 8 8 8 8 8
ro.20
5.0 5.0
8 4
\0.18 0.43
5.0
12
0.70 0.22 0.64 0.70 0.45 0.50 JO. 37 \0.36 0.11
Reduced level2
Gain, gms./day
No. birds
0.78
11.0 5.0
12 4
—
—
—
4.5 4.0
8 4
—
—
0.82 0.72 0.42 0.79 0.36 0.28 0.10 0.27 0.10
—
8.0 5.0 4.0 4.0 4.0
4 4 8 4 16
1
Essential amino acid levels in ration 1 were as follows (in percent): arginine, 1.28; histidine, 0.43; lysine, 1.15; leucine, 1.30; isoleucine, 0.80; valine, 0.95; phenylalanine, 0.67; tyrosine, 0.66; tryptophan, 0.20; methionine, 0.38; cystine, 0.35; and threonine, 0.78. The D-isomer of isoleucine, valine, and threonine was considered to be inactive. 2 Based on values given by March el al. (1962).
White Plymouth mating. The colored chicks were produced by mating White Plymouth Rock females with colored males produced from a Red Vantress X White Plymouth Rock mating. A variety of colors and color patterns was found among the colored chicks. The chicks were not sexed. The chicks were reared in electricallyheated batteries with wire floors. Feed and water were supplied ad libitum. Chicks were fed a practical type broiler ration for one day after being taken from the incubator and then distributed among experimental groups on the basis of body weight and color. Experimental rations were fed for approximately two weeks and the birds were then returned to the practical ration. Pictures were taken about one week later to allow some of the modified feathers to develop to a point where the modifications were visible. RESULTS AND DISCUSSION
The mean weight gains are summarized in Table 2. Weight gains were reduced from 11.0 to from 4.0 to 5.5 grams per day by modifying ration 1 and 2. Herring meal
is an excellent source of lysine, so the mean weight gain was not reduced as much when this amino acid was removed from the amino acid mixture used with ration 2. Large numbers of chicks were not fed each ration in these tests. Most of the changes in feather structure noted were evident after the first experiment. Lysine, threonine, and methionine and cystine deficient rations were fed in our studies on peanut meal (Anderson and Warnick, 1965) and cottonseed meal (Anderson and Warnick, 1966), so it did not seem justified to feed additional chicks deficient rations. The observations made in this report are based primarily on the experiments conducted with rations 1 and 2. Some statements, however, necessarily also draw upon observations made during the above listed experiments. Abnormal feather development was noted when arginine, isoleucine, leucine, valine, or phenylalanine and tyrosine were deficient in the diet. Chicks fed the tryptophan deficient diet showed some ragged feathering but not to the degree found among chicks fed the diets with the five
AMINO ACID DEFICIENCY SYMPTOMS
859
FIG. 1. Chick on the right was fed ration 1, while the chick on the left was fed ration 1 with the valine level reduced to 0.5 percent.
deficiencies listed first. When the ration was deficient in histidine, lysine, threonine, or methionine and cystine, growth rate and feather development were slower than with the basic ration, but the chicks appeared normal otherwise. Even the achromatosis usually associated with lysine deficiency did not appear when part of the lysine was removed from rations 1 and 2. This lack of pigment had been noted several times when other lysine deficient rations were fed in our laboratory. Abnormal feather development had not come to our attention in our earlier experiments in which diets deficient in histidine, threonine, or methionine and cystine were fed. However, abnormal feathering may develop if rations containing more serious deficiencies of these amino acids are fed. The abnormal feather conditions noted among chicks fed rations deficient in arginine, valine, leucine, isoleucine, tryptophan, or phenylalanine and tyrosine were somewhat similar. If there was a difference between any two deficiencies it was only in the degree to which the abnormalities de-
veloped. Similar symptoms were produced when either ration 1 or 2 was made deficient in one of the amino acids. The abnormalities noted included the ragged appearance reported by Hegsted et al. (1941) and the spoon-shaped and curled feathers noted by Sanders et al. (1950). Often the rachis of each secondary was found to curl toward the body when the wing was held in the open position. The small tracts of feathers on the shoulders curled upwards; these changes were noted within a few days after the chicks were started on the deficient rations. Many feathers lacked their characteristic smooth surface. When these feathers were examined under the microscope it was found that the barbules on adjacent barbs did not form the normal interlocking structure, and the barbules did not spread as much as in normal feathers. Hooklets were usually present although, in some cases, the barbules appeared to have been broken off with a loss of the hooklets. These conditions gave the feathers a ragged and sometimes thin appearance.
860
J. O. ANDERSON AND R. E. WARNICK
FIC. 2. Chick fed ration 2 with the arginine removed from the amino acid mixture.
The chick on the right in Figure 1 was fed ration 1 during the experimental period. Chicks fed this ration were not as smoothly feathered as chicks fed a well balanced broiler ration, but their feathering was smoother than that of chicks fed any of the six deficient rations listed above. The
FIG. 3. Chick fed ration 1 with the leucine level reduced to 0.7 percent.
chick on the left in Figure 1 was fed ration 1 with the valine level reduced from 0.95 to 0.5 percent. This chick was typical of the valine deficient birds. The feathering was ragged and the secondaries showed a marked curl. When the wing was held in the normal position the secondaries extended above the bird's backline. The chick in Figure 2 was fed ration 2 with the arginine removed from the amino acid mixture. Fewer than 25 percent of the birds fed this ration were affected to this extent. The feather condition was similar to that described by Hegsted et al. (1941) and Sanders et al. (1950). Some of the arginine deficient birds were unsteady while they were standing. Weak birds were noted in groups of chicks fed rations deficient in other amino acids, but the unsteadiness while standing and the unusual gait seemed to be more common or even peculiar to the arginine deficient chicks. Some birds fed this ration appeared normal except for the smaller body size. The chick in Figure 3 was fed ration 1 and the leucine level reduced from 1.3 to 0.7 percent. The feathering of this chick was affected to a greater degree than that of the average bird fed the leucine deficient rations. The leucine deficient chicks showed
FIG. 4. Chick fed ration 1 with the isoleucine level reduced to 0.45 percent.
AMINO ACID DEFICIENCY SYMPTOMS
more ragged, curled, and twisted feathers. Even the primary and secondary coverts and the breast feathers of the pictured chick were curled abnormally. The chick in Figure 4 was fed ration 1 wtih isoleucine level reduced from 0.8 to 0.4S percent. The feathering of birds fed this ration was less affected than that of the birds fed the valine, arginine, or leucine deficient diets. About one-third of these birds showed flight feathers with a ragged and thin appearance. Colored chicks fed the rations deficient in phenylalanine and tyrosine showed changes in the feather pigments (see Figure 5). The most obvious difference was noted with black feathered chicks. These chicks showed a gray bar across the flight feathers; the color of body feathers was also affected. Red birds showed some less obvious loss of pigments. Both colored and white chicks showed ragged feathering. SUMMARY
Chicks were fed rations deficient in each of the essential amino acids for two weeks. The deficient rations were produced by removing in turn approximately 46 percent of each essential amino acid from rations with a well balanced mixture of the essential amino acids. Reducing the essential amino acid levels reduced the mean growth
FIG. 5. Chick fed ration 2 without the phenylalanine and tyrosine of the amino acid mixture.
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rate from 11 to approximately 4.S grams per day. Abnormal feather structures were evident when the ration was deficient in leucine, arginine, valine, isoleucine, tryptophan, or phenylalanine and tyrosine. Abnormalities included curled feathers, spoonshaped feathers, and feathers with a ragged and thin appearance. The feather abnormalities produced with each of the six listed deficiencies were somewhat similar, but these symptoms developed to different degrees. Reduced feather pigmentation was noted when a phenylalanine and typrosine deficient ration was fed to colored chicks. REFERENCES Anderson, J. O., and R. E. Warnick, 1965. Amino acid deficiencies in peanut meal and in corn and peanut meal rations. Poultry Sci. 44: 10661072. Anderson J. O., and R. E. Warnick, 1966. Sequence in which essential amino acids become limiting for growth of chicks fed rations containing cottonseed meal. Poultry Sci. 45: 8489. Dobson, D. C , J. O. Anderson and R. E. Warnick, 1964. A determination of the essential amino acid proportions needed to allow rapid growth in chicks. J. Nutrition, 82: 67-75. Fritz, J. C , J. H. Hooper, J. L. Halpin and H. P. Moore, 1946. Failure of feather pigmentation in bronze poults due to lysine deficiency. J. Nutrition, 3 1 : 387-396. Grau, C. R., 1945. Deformity of the tongue associated with amino acid deficiencies in the chick. Proc. Soc. Exp. Biol. Med. 59: 177-178. Hegsted, D. M., G. M. Briggs, C. A. Elvehjem and E. B. Hart, 1941. The role of arginine and glycine in chick nutrition. J. Biol. Chem. 140: 191-200. Klain, G. J., D. C. Hill, J. A. Gray and H. D. Branion, 1957. Achromatosis in the feathers of chicks fed lysine-deficient diets. J. Nutrition, 61 : 317-328. Klain, G. J., H. M. Scott and B. C. Johnson, 1960. The amino acid requirements of the growing chick fed a crystalline amino acid diet. Poultry Sci. 39 : 39-44. March, B. E., J. Biely and H. L. A. Tarr, 1962. Nutrient composition of British Columbia herring meal. Fisheries Research Board of Canada Circular No. 26.
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J. O. ANDERSON AND R. E.
Ousterhout, L. E., 1960. Survival time and biochemical changes in chicks fed diets lacking different essential amino acids. J. Nutrition, 70: 226-234. Sanders, B. G., S. O. Brown and J. R. Couch, 1950. A feathering syndrome in chicks after feeding optimal levels of lysine in the absence
WAENICK
of arginine. Proc. Soc. Exp. Biol. Med. 74: 114— 117. Wilkening, M. C , B. S. Schweigert, P. B. Pearson and R. M. Sherwood, 1947. Studies on the requirement of the chick for tryptophan. J. Nutrition, 34: 701-714.
The Utilization of Phytate Phosphorus by Poultry—A Review T. S. N E L S O N
Research and Development Division, International Minerals and Chemical Libertyville, Illinois 60048
Corporation,
(Received for publication November 8, 1966)
T
HE metabolism of phosphorus derived from plant tissues is one of the least understood and most debated subjects in the field of mineral nutrition. A small proportion of the total phosphorus in plants is inorganic phosphates located primarily in the vegetative tissues. Most of the phosphorus is in a variety of organic compounds found primarily in the seed, and phytate phosphorus is the predominant organic form. The amount of phytate phosphorus utilized by animals has economic importance because seeds are the major plant-source ingredients used in feeds. Many scientists currently use the rule that monogastric animals metabolize only one-third of the phosphorus in plant materials. This assumption is based on the report by the Committee on Animal Nutrition, National Academy of Sciences-National Research Council (NAS-NRC) (1960) that approximately 30% of the phosphorus in plant materials in non-phytate and can be considered to be utilized by animals. Many scientists using a variety of experimental procedures have studied the ability of different species of animals to utilize phytate phosphorus. However, they have failed to define to everyone's satisfaction
the extent of phytate phosphorus availability in strictly quantitative terms. Certain authors have reported it was utilized to a limited extent. Others have considered it was highly available to animals. Kastelic and Forbes (1961) and Taylor (1965) reviewed this subject. They stressed that there is still no general agreement on the extent to which different species of animals at various ages utilize phytate phosphorus. Studies of Phytate Phosphorus Utilization Heuser et al. (1943), McGinnis et al. (1944), Singsen et al. (1947), Gillis et al. (1949), and Sunde and Bird (1956) observed that natural phytate was a poor source of phosphorus for various species of poultry. Conversely, Sieburth et al. (1952) reported the phosphorus in finely ground whole wheat flour was almost completely available to chicks for growth but was less available than inorganic phosphate for bone deposition. Temperton et al. (1965a, b, c) concluded that pullet chicks less than four weeks old, growing pullets reared to 18 weeks of age and laying hens were able to achieve effective utilization of organic sources of phosphorus for growth and bone formation. Several investigators have fed various