862 J. O. ANDERSON AND R. E.
WAENICK
Ousterhout, L. E., 1960. Survival time and bio-
chemical changes in chicks fed diets lacking dif-
ferent 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
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
(Received for publication November 8, 1966)
T HE metabolism of phosphorus de-
rived 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 pri-
marily in the vegetative tissues. Most of
the phosphorus is in a variety of or-
ganic compounds found primarily in the
seed, and phytate phosphorus is the pre-
dominant 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 Nutri-
tion, National Academy of Sciences-Na-
tional Research Council (NAS-NRC)
(1960) that approximately 30% of the
phosphorus in plant materials in non-phy-
tate and can be considered to be utilized by
animals.
Many scientists using a variety of exper-
imental procedures have studied the ability
of different species of animals to utilize
phytate phosphorus. However, they have
failed to define to everyone's satisfaction
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 re-
quirement of the chick for tryptophan. J. Nu-
trition, 34: 701-714.
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Corporation,
the extent of phytate phosphorus availabil-
ity 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) re-
viewed 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) ob-
served 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
 UTILIZATION OF PHYTATE PHOSPHORUS
phytates as the isolated but impure chemi-
cal compound rather than relying on feed
ingredients as the source. Lowe et al.
(1939) reported that chicks did not
efficiently utilize phytate phosphorus iso-
lated from wheat bran. Singsen and Mit-
chell (1945) and Matterson et al. (1946)
found calcium magnesium phytate was a
poor source of phosphorus for the turkey
poult. Gillis et al. (1948) showed that
chicks were unable to utilize relatively pure
calcium phytate. Waldroup et al. (1964a)
reported the availability of the phosphorus
in calcium phytate and sodium phytate was
less than that of sodium phosphate. Harms
et al. (1962) and Waldroup et al. (1964a)
concluded that the phosphorus in phytic
acid was highly available to the chick.
Few investigators have studied the quan-
titative utilization of phytate phosphorus
by poultry. Gillis et al. (1953) reported
White Leghorn hens utilized phytate phos-
phorus approximately one-half as effective-
ly as the phosphorus in dicalcium phos-
phate. Gillis et al. (1957) fed chicks and
turkeys P 32 labeled calcium phytate and P 32
labeled monosodium ortho phosphate and
then measured the amount of radioactivity
retained in the tibia. They concluded that
chicks utilized the phosphorus in calcium
phytate only 10% as effectively as that in
monosodium ortho phosphate and that the
relative utilization of calcium phytate
phosphorus by the turkey was less than 2%.
The availability of phytate phosphorus
has also been studied in phosphorus bal-
ance trials. Nikolaiczuk (1950) reported
that chicks retained 14% to 18% of the
phytate phosphorus in the dietary ingredi-
ents. Not all of the phytate phosphorus was
recovered in the feces and the author con-
cluded that degradation processes other
than hydrolysis accounted for this unrecov-
ered phytate. Ashton et al. (1960) fed P 32
labeled calcium phytate and observed that
chicks four-weeks old retained approxi-
863
mately 20% of the phytate phosphorus
whereas six-week old chicks retained 36%
to 49% of this phosphorus. They con-
cluded the chicks utilized one-fifth of the
phytate phosphorus. Temperton and Cassi-
dy (1964)reported chicks retained approxi-
mately 60% of the phytate phosphorus and
only 50% of the non-phytate phosphorus.
They indicated that the chick's total need
for phosphorus could be met with plant-
source phosphorus.
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It is apparent that wide disagreement
has existed between investigators on the
ability of the chick to utilize phytate phos-
phorus. Varied experimental methods and
materials were used and may have con-
tributed to this disagreement. These varia-
bles included the source of phytate phos-
phorus, criteria of response, age of the test
animals, and calcium and vitamin D 3 lev-
els in the experimental diets.
Sources of Phytate Tested
The sources of the phytates tested may
have caused some of the discrepancies be-
tween reports on phytate phosphorus utili-
zation by poultry. Phytate phosphorus oc-
curs in plants as the mixed calcium-magne-
sium-potassium salt of phytic acid (Ander-
son, 1915c; Averill and King, 1926). The
availability of this "natural" phytate phos-
phorus may differ from that in a chemically
isolated fraction for at least two reasons.
The first is the fact that phytic acid-protein
complexes are formed in certain types of
processed seed meals that reduce the solu-
bility of these proteins (Fontaine et al.,
1946). Such complexes may also cause fur-
ther reduction in the availability of the
phytate phosphorus by delaying the onset
of enzymatic attack during the digestive
process. Another reason is the source of the
natural phytate phosphorus may influence
its availability to the animal. Some feed
ingredients contain the enzyme phytase
which hydrolyzes phytate to phosphoric
864 T. S. NELSON
acid (Anderson, 1915a, b). Wheat, rye and
barley are high in phytase activity whereas
oats, maize and various seed meals contain
little or none of the enzyme (McCance
and Widdowson, 1944; Mollgaard, 1946).
Therefore, phytate phosphorus should be
more available in ingredients containing
this enzyme or in diets containing ingredi-
ents with a high phytase content.
Chemical forms of phytate appear to
offer both advantages and disadvantages
for phosphorus assay purposes. These com-
pounds are not bound as the protein com-
plex and presumably should give a more
reliable estimate of phytate phosphorus
utilization per se. It would appear that
compounds such as magnesium phytate are
the most desirable compounds to test since
they are similar in composition to natural
phytate. However, neither is the same com-
pound that existed in the plant.
Calcium phytate and calcium magnesium
phytate were tested in the work cited
above. They were found to be a poor
source of phosphorus, which suggested that
natural phytate was also poorly utilized.
Sodium phytate and phytic acid were other
chemical phytates tested in the work cited
above. Both were reported to be better
sources of phosphorus than calcium phy-
tate. However, neither was the form found
in the plant. Therefore, for reasons men-
tioned below, they cannot be used to deter-
mine phytate phosphorus utilization.
The purity and solubility of any phytate
compound may influence the apparent
availability of the phosphorus. Analyses in
this laboratory have shown that calcium
phytate, sodium phytate, and phytic acid
were contaminated with ortho phosphate.
This form of phosphorus would be metab-
olized by the animal and could lead to er-
rors in estimating the utilization of phytate
phosphorus. Calcium phytate is insoluble
and is relatively inert to acid or basic hy-
drolysis in the pH ranges that exist in the
digestive tract of poultry. Conversely, so-
dium phytate and phytic acid are more sol-
uble and should be partially hydrolyzed to
phosphoric acid which can be utilized by
the chick. Accordingly, one cannot extrapo-
late the availability of the phosphorus in
sodium phytate or phytic acid to naturally
occurring phytate phosphorus.
Criteria of Response
Percent bone ash and weight gain have
been the most common criteria used to
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study the utilization of phytate phos-
phorus. P 32 deposition from labeled phytate
and phosphorus balance studies have been
used to a lesser extent.
The percent of bone ash is one of the
most sensitive practical criteria for evaluat-
ing the availability of dietary phosphorus.
It is more accurate than body weight (Nel-
son and Walker, 1964; Dilworth, 1966)
and is little affected by other dietary varia-
bles that influence growth.
Body weight gain is not an accurate
measure of phosphorus utilization and its
use has led to misleading conclusions. Sing-
sen et al. (1947) and Sieburth et al.
(1952) reported phytate phosphorus was
more available for growth than for bone
calcification. Vanderpopuliere et al. (1961)
concluded plant-source phosphorus was
readily available to support growth but
that it would not increase the percent bone
ash when the calcium:phosphorus ratio was
narrowed from 4:1 to 1:1. Waldroup et al.
(1964a), in an opposing view, reported cal-
cium phytate phosphorus was more avail-
able for bone deposition than for growth. A
more realistic view was expressed by Tay-
lor (1965), commenting on the theory that
the animal can partition phytate phospho-
rus between growth and bone calcification:
"If it is agreed that the P of phytate is in
all probability absorbed as phosphate ions,
the concept of a greater or lesser 'availabil-
ity' for a particular physiological function
becomes quite unacceptable."
Nelson et al. (1965) published data sup-
 UTILIZATION OF PHYTATE PHOSPHORUS
porting this statement and explaining the
differences observed between growth and
bone deposition. Chicks were fed purified
diets practically devoid of calcium and
phosphorus. These diets were supplemented
with inorganic sources of calcium and
phosphorus at ratios of 4:1 and 2:1. The
chicks gained weight when the calcium con-
tent of these diets was reduced to narrow
the ratio. However, the source of supple-
mental phosphorus determined whether or
not the percent bone ash increased as the
calcium content of the diet was decreased.
These data, obtained in the absence of phy-
tate phosphorus, were similiar to those of
Singsen et al. (1947), Sieburth et al.
(1952) and Vanderpopuliere et al. (1961)
who reported the possible partition of phy-
tate phosphorus between growth and bone
deposition. It is probable that the improved
weight gain they observed was a response
to a more favorable calcium level rather
than a peculiar partition in the utilization
of phytate phosphorus. If this assumption
is true, then the availability of phytate
phosphorus in finely ground whole wheat
flour was less than indicated by Sieburth et
al. (1952).
Common (1939) reported the phytate
was inactive in the body of the chick and
the poult and passed out in the feces as the
intact molecule. Singsen et al. (1950) con-
cluded the phosphorus in the phytate mole-
cule moved quickly and easily throughout
the body of the turkey poult. Although
these authors disagreed on the metabolism
of the phytate molecule, they did agree on
the probable availability of the phosphorus.
Singsen's group fed poults P 32 labeled cal-
cium phytate and found radioactivity in
the blood, down and muscle tissue within
one hour after dosing. They stated it was
unlikely that the low phytase activity in
the intestine would hydrolyze P 32 from the
phytate molecule so rapidly. Although P32
was deposited in the bone salts, the un-
labeled phytate phosphorus did not support
865
bone calcification. They explained that the
exchange reaction probably was the mech-
anism observed, and if true, the phytate
molecule removed as much phosphorus from
the body as it brought in. This agreed with
Common's belief that the intact phytate
molecule was excreted in the feces.
The exchange reaction is the simple
chemical exchange of the same element be-
tween two different compounds. The ex-
change of P 32 from phytate with P from an
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inorganic source would render the P 32 avail-
able. It would appear, based on radioactivi-
ty measurements, that phytate phosphorus
is utilized. However, if it is replaced with
inorganic P which is rendered unavailable
when incorporated into the phytate mole-
cule, there is no net utilization of phytate
phosphorus.
Gillis et al. (1957) used P 32 labeled cal-
cium phytate and P 32 labeled monosodium
ortho phosphate to study phytate phospho-
rus utilization. Their conclusions, discussed
above, were based on the amount of P 32
retained in the tibia. They also demon-
strated that P 32 from NaH 2 P 32 0 4 exchanged
readily with the phosphorus from calcium
phytate in vitro. This agreed with the
theory Singsen et al. (1950) used to explain
the apparent utilization of phytate phos-
phorus by poults. The work of Gillis et al.
(1957) emphasized the fact that the recov-
ery of radioactivity in the body does not
necessarily represent a net utilization of
phytate phosphorus. It is possible for the
net utilization to be zero because of the
exchange reaction.
The effect of age on the ability of poul-
try to utilize phytate phosphorus has not
been clearly defined. The data available
suggest that the utilization of phytate
phosphorus increases with increasing age to
maturity. The degree to which this is true,
however, has not been established, primarily
because of the types of experimental diets
used.
McGinnis et al. (1944) compared the
866 T. S. NELSON

utilization of the phosphorus in a natural equal amounts of calcium and phosphorus.

diet by chicks to four and eight weeks of Plant phytate is the mixed calcium magne-
age. The chicks had a higher percentage of sium potassium salt of phytic acid and
bone ash at eight weeks. This could have therefore contains less calcium than phos-
been the result of either increased utiliza- phorus. The availability of phytate calcium
tion of phytate phosphorus or a reduced has not been established and it is presum-
phosphorus requirement. Gillis et al. ably possible, based on the work cited
(1953, 1957) observed differences in the above, that little of it is available. Further-
ability of the chick and the hen to utilize more, it is also presumably possible for the
phytate phosphorus. The hen diet con- phytate molecule to inactivate a portion of

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tained wheat bran, which probably con- the supplemental calcium in the digestive
tained the enzyme phytase. Conceivably, tract. Theoretically, this could occur be-
part or all of the increased ability of the cause of the low calcium-phosphorus ratio
hen to utilize phytate phosphorus could of plant phytate compared to calcium phy-
have been due to wheat bran phytase. tate.
Ashton et al. (1960) found differences in Another role of calcium on apparent
phytate phosphorus retention of four-week availability of phytate phosphorus concerns
and six-week old chicks fed the same diets. the dietary levels fed in relation to the
Temperton et al. (1965a, b, c) reported available phosphorus. This was discussed
that pullets and laying hens could utilize above, but it should be reemphasized that
organic sources of phosphorus. However, at it is fallacy to conclude that responses ob-
least three ingredients in their diets proba- tained by changing dietary calcium levels
bly contained high phytase activity. result from changes in the availability of
Therefore, while older poultry may utilize phytate phosphorus.
phytate phosphorus more efficiently than
younger birds, the use of feed ingredients Vitamin D
that contained phytase activity in some ex- The effect of vitamin D on phosphorus
periments may have overemphasized this utilization has received renewed attention
age difference. in recent years. The primary function of
this vitamin, demonstrated by Nicolaysen
Calcium. et al. (1953) and by Keane et al. (1956) is
Calcium has been shown to have an ad- to promote calcium absorption. This pro-
verse effect on the availibility of phytate cess is an active one which proceeds against
phosphorus. Conversely, the phytate mole- an electrochemical gradient (Wasserman et
cule has been observed to reduce the avail- al, 1961). Nicolaysen et al. (1953) con-
ability of calcium. Phytate supplied either cluded that the increased phosphorus
by dietary ingredients or in a chemical absorption they observed was secondary to
form reduced calcium absorption from the the calcium absorption. Their work re-
intestine of humans (McCance and Wid- ceived the support of Harrison and Harri-
dowson, 1942a, b; Krebs and Mellanby, son (1961) who concluded that phosphate
1943; Bronner et al, (1954). Hoff-Jorgen- transport was dependent of calcium. Ac-
sen (1946) and Mellanby (1949) observed cording to Rasmussen and DeLuca (1963),
a similar effect in dogs. Melanby attributed the only regulation of phosphate absorption
this to the precipitation of insoluble penta- by vitamin D is indirectly through changes
calcium phytate. in calcium absorption.
Calcium phytate contains approximately Reports have been published suggesting
 UTILIZATION OF PHYTATE PHOSPHORUS 867

that vitamin D 3 improved the utilization of
phytate phosphorus. Lowe et al. (1939),
Singsen and Mitchell (194S) and Singsen
et al. (1947) increased the bone ash in
chicks and poults by feeding higher levels
of vitamin D 3 . McGinnis et al. (1944) re-
ported chicks responded to increased levels
of vitamin D 3 in a manner suggesting bet-
ter utilization of phytate phosphorus. Gillis
et al. (1949) observed that high levels of
ferentiate between the response to vita-
min D 3 because of the imbalanced calcium-
available phosphorus ratio and in enhanc-
ing phytate phosphorus utilization. With
both of these actions occurring simulta-
neously, the results could overemphasize
the value of vitamin D 3 for improving phy-
tate phosphorus utilization.
A second theory suggested that vitamin
D increased the production of phytase in

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vitamin D 3 improved the performance of the intestine or that it enhanced the activi-
chicks fed either calcium phytate or natu- ty of the phytase contained in the feed in-
ral phytate phosphorus. Gillis et al. (1957) gredients. The report by Spitzer et al.
found that vitamin D 3 caused a small in- (1948) does not agree with this theory.
crease in the retention of phytate P 32 in the They were unable to increase the intestinal
bones. Waldroup et al. (1964b) increased phytase activity by adding vitamin D to
the weights of the chicks fed either calcium tissue homogenates. Pileggi et al. (1955)
phytate or dicalcium phosphate by increas- published evidence to support this theory.
ing the vitamin D 3 content of the diet. They reported that vitamin D increased
At least two theories have been put forth the intestinal phytase activity two to five
to explain the mode of action of vitamin times in the intact animal. Roberts and
D 3 . Singsen et al. (1950) suggested that Yudkin (1961) also found that this vita-
phytate phosphorus was labile, but in the min increased phytase activity in rats. How-
absence of this vitamin the phytate mole- ever, this theory has not been investigated
cule probably removed as much phosphorus fully to determine its merits.
from the body as it brought in. However,
under the influence of vitamin D 3 another Phytase Enzyme
radical replaced part of the phytate phos- The presence of the enzyme phytase in
phorus which then became available to the certain feed ingredients and its possible se-
animal. The improved phosphorus utiliza- cretion in the intestine were discussed ear-
tion in the presence of vitamin D 3 repre- lier in this review. Additional information
sented this released phytate phosphorus, on the action of phytase was reported by
rather than the increased utilization of the Courtois (1945) and by Courtois and Mas-
non-phytate phosphorus. son (1950). Plant phytase was specific for
The effect of vitamin D 3 in increasing phytate phosphorus and cleaved phosphate
the utilization of the inorganic phosphorus from alternate rather than adjacent carbon
cannot be totally discounted, however. It is atoms. Sandegren (1948) and Mattson and
well documented that, at an optimum calci- Koutler-Andersson (1947) indicated that
um to available phosphorus ratio, little, if this enzyme was active in a narrow pH
any, response is obtained from vitamin D 3 . range. Proteins precipitated phytate on the
Conversely, a response can be obtained if acid side, but it was precipitated by calci-
the ratio is unbalanced. Vitamin D 3 re- um and magnesium ions in an alkaline con-
sponses were obtained in most of the exper- dition. Plant phytase can hydrolyze phy-
iments cited above with diets containing tate phosphorus, and the conditions for its
imbalanced levels of calcium and available action suggest that the enzyme would be
phosphorus. It was impossible to dif- active in the digestive tract of animals.
868 T. S. NELSON

Singsen and Mitchell (1944) studied the middlings. These two ingredients supposed-
effect of feeding an ingredient with an ap- ly contained phytase activity. The dif-
parent high phytase activity. The addition ference between their results and those
of sun-cured alfalfa meal, which presum- of Singsen and Mitchell (1944) and Tem-
ably contained phytase activity, to a chick perton et al. (1965a, b, c) suggests that
diet promoted weight gain and bone ash as plant-source feed ingredients either vary in
well as supplemental inorganic phosphate their original phytase content or that the
whereas dehydrated alfalfa meal was enzyme was labile under the handling con-
ineffective. They concluded that the sun- ditions of the feed ingredients. It appears
cured sample contained phytase activity that specific feed ingredients cannot be

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that was apparently destroyed in the dehy- considered to be a consistent source of phy-
drated sample. If their conclusion was cor- tase.
rect, then it appears that the enzyme also Curtois and Valentino (1947) found that
hydrolyzed the phytate phosphorus con- purified phytase, extracted from wheat
tained in the other feed ingredients. Tem- bran, was ineffective in the rat. The
perton et al. (1965a, b, c) presented indi- difference between their results and those
rect evidence that plant phytase was effec- of Warden and Schaible (1962) may have
tive in feeds. Their diets contained 32 to reflected differences in phytase enzymes
36% wheat and 10% barley meal which from plant and bacterial sources.
have been reported to be sources of phy-
tase. These diets also contained three per- Plant Phosphorus
cent grass meal which may have been Common (1940) reported the phytic
similar to the sun-cured alfalfa meal acid phosphorus content of feedstuffs de-
studied by Singsen and Mitchell (1944). rived from processed oil seeds, including
Warden and Schaible (1962) appear to soya bean meal, was one-half to two-thirds
have established that bacterial phytase can of the total phosphorus. Phytic acid phos-
be active in the digestive tract. They found phorus accounted for two-thirds to three-
the addition of lysed E. coli cellular mate- fourths of the total phosphorus in cereal
rial to the diet improved both growth and grains.
bone development and suggested this was a
Mollgaard (1946) showed slightly
response to phytase or similar enzymes.
higher concentrations of phytate phospho-
It appears, in view of these reports, that rus in processed oil seeds (70-80%) and
some of the differences observed in the in cereal grains (65-85%).
ability of the various species of poultry to The non-phytate phosphorus content of
utilize phytate phosphorus can be ex- those ingredients mentioned above ranged
plained by the presence or absence of phy- from 15-50%. The degree to which this is
tase in the experimental diets. utilized by the animal has not been clearly
Although the work cited above suggests established. It is questionable, from the
phytase activity can be a determining fac- data available, whether any portion of the
tor in phytate phosphorus utilization, the phytate phosphorus should be considered
conditions for its activity have not been available. Rations formulated with one part
clearly established. McGinnis et al. (1944) plant protein source for each two parts
found chicks made inefficient use of phy- plant energy source will contain approxi-
tate phosphorus in a diet that contained mately 30% non-phytate phosphorus. Con-
20% wheat bran and 10% wheat flour sidering this to be available, phosphorus
 UTILIZATION OF PHYTATE PHOSPHORUS
would agree with the NAS-NRC recom-
mendation that 30% of the plant phospho-
rus is available to the animal.
The NAS-NRC recommendation does
not consider (a) variations in the phytate
content of different ingredients, (b) the
availability of the non-phytate phosphorus,
and (c) the possible utilization of some of
the phytate phosphorus. In view of the fact
that animals can utilize some portion of the
plant phosphorus, and based upon the dis-
cussion above, the NAS-NRC recom-
mendation appears to be a close estimate of
the available phosphorus in a mixture of
plant materials but not for individual in-
gredients.
Conclusions
The preponderance of data suggests that
phosphorus is absorbed as the inorganic
phosphate ion. Therefore, the ability of the
various species of poultry to utilize phytin
phosphorus will depend, in the simplest
terms, on their ability to liberate phosphate
ions from the phytate molecule by hydroly-
sis. There is no evidence showing unques-
tionable proof that phytate is absorbed and
utilized intact by any animal species. Pres-
ent knowledge of calcium and phosphorus
utilization and the role of vitamin D 3 per-
mits tentative conclusions on the various
species of poultry to utilize phytin phos-
phorus.
1. In order to be utilized, phytin phos-
phorus must be hydrolyzed to yield inor-
ganic phosphate.
2. The chick absorbs and utilizes dietary
phosphorus without regard to its organic or
inorganic origin. The chick cannot parti-
tion the phosphorus derived from the phy-
tate molecule between different metabolic
functions in a manner different from that
derived from inorganic sources.
3. Young birds have very limited ability
to hydrolyze phytin phosphorus. This abili-
869
ty increases with age up to maturity; how-
ever, the magnitude of this increase has not
been accurately quantified.
4. Calcium, at levels required by poul-
try, has an adverse effect upon the utiliza-
tion of phytin phosphorus. The availability
of phytate calcium has not been estab-
lished.
5. Vitamin D 3 probably enhances the
utilization of phytate phosphorus by ani-
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mals to a limited extent. However, the
magnitude of this increase is small and ap-
pears to be dependent upon several other
dietary interrelationships.
6. Phytase enzyme, present in certain
feed ingredients and possibly secreted by
the intestine, hydrolizes phytate phospho-
rus to a form that can be utilized. Under
certain conditions this enzyme will hydro-
lyze dietary phytate phosphorus making it
available to the animal. This appears to be
the most effective force at work in the di-
gestive tract that results in utilizaion of
phytate phosphorus.
7. The National Academy of Sciences-
National Research Council's recommenda-
tion that 30% of the total plant phospho-
rus is available to animals can usually be
applied to a mixture of ingredients but not
to individual ingredients.
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Anderson, R. J., 1915a. The hydrolysis of phytin
by the enzyme phytase contained in wheat bran.
J. Biol. Chem. 20:475-482.
Anderson, R. J., 1915b. The hydrolysis of the or-
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the enzyme phytase. J. Biol. Chem. 20: 483-
491.
Anderson, R. J., 1915c. Concerning phytin in
wheat bran. J. Biol. Chem. 20: 493-500.
Ashton, W. M., C. Evans and P. C. Williams,
1960. Phosphorus compounds of oats. II. The
utilisation of phytate phosphorus by growing
chicks. J. Sci. Food Agric. 11: 722-727.
Averill, H. P., and C. G. King, 1926. The phytin
content of foodstuffs. J. Am. Chem. Soc. 48:
724-728.
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Bronner, F., R. S. Harris, C. J. Maletskos and C.
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Common, R. H., 1939. Phytic acid in mineral me-
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Courtois, J., 1945. Phytase. I. Comparative action
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Courtois, J., and M. Masson, 1950. Phytase XVI.
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