Review: Anti-nutritional effects of phytic acid in diets


for pigs and poultry current knowledge and directions
for future research
T. A. Woyengo and C. M. Nyachoti1
Department of Animal Science, University of Manitoba, Winnipeg, Manitoba, Canada R3T 2N2.
Received 13 February 2012, accepted 23 September 2012.


Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.230.235 on 05/17/20

Woyengo, T. A. and Nyachoti, C. M. 2013. Review: Anti-nutritional effects of phytic acid in diets for pigs and poultry
current knowledge and directions for future research. Can. J. Anim. Sci. 93: 921. Plant feedstuffs contain phytic acid (PA),
which is a storage form of phosphorus. Phytic acid is, however, poorly hydrolyzed by pigs and poultry, and it has a
capacity to complex dietary nutrients, thereby reducing nutrient digestibility. Reduced nutrient digestibility by PA implies
reduced efficiency of utilization of the nutrients and increased discharge of the unabsorbed nutrients to the environment.
Phytic acid has also recently been shown to increase the endogenous nutrient losses (ENL) in pigs and poultry. Because the
increased ENL in the gastrointestinal tract are associated with increased maintenance requirement for the lost nutrients
and of energy spent on their secretion, an increase in ENL due to PA implies that there are other adverse effects of PA on
nutrient utilization in addition to reducing nutrient digestibility. In this review, the effects of PA on performance of pigs
and poultry, and on the digestibility and ENL in these animals are discussed in detail. Also, the mechanisms by which PA
reduces nutrient digestibility and increases gastrointestinal ENL in pigs and poultry are discussed, and areas that need
further research to gain more insight into these mechanisms are suggested.

Key words: Phytic acid, pig, poultry, nutrient utilization

Woyengo, T. A. et Nyachoti, C. M. 2013. Effets anti-nutritionnels de l’acide phytique dans la ration des porcs et de la

For personal use only.

volaille connaissances actuelles et pistes pour les recherches à venir. Can. J. Anim. Sci. 93: 921. Les aliments végétaux
renferment de l’acide phytique (AP), composé servant à stocker le phosphore. Malheureusement, les porcs et la volaille
hydrolysent mal cet acide, qui, de plus, a tendance à rendre les nutriments plus complexes, donc à en réduire la digestibilité.
Une digestibilité plus faible des éléments nutritifs suppose en retour une assimilation moins efficace et l’expulsion dans
l’environnement d’une plus grande quantité de nutriments non absorbés. Récemment, on a aussi montré que l’AP
augmente les pertes endogènes de nutriments (PEN) chez le porc et la volaille. L’augmentation des PEN dans le tube
digestif est associée aux besoins de maintenance plus élevés nécessaires pour compenser les nutriments perdus et l’énergie
dépensée lors de leur sécrétion. La hausse des PEN attribuable à l’AP laisse présumer que celui-ci a d’autres effets
secondaires sur l’assimilation des nutriments qu’une digestibilité réduite. L’analyse examine en détail les conséquences de
l’AP sur le rendement des porcs et de la volaille et sur la digestibilité ainsi que les PEN chez ces animaux. On y discute aussi
des mécanismes par lesquels l’AP réduit la digestibilité des nutriments et augmente les PEN dans le tube digestif des porcs
et de la volaille. Les auteurs préconisent qu’on approfondissent les recherches dans ces domaines afin d’en apprendre
davantage sur ces mécanismes.

Mots clés: Acide phytique, porc, volaille, assimilation des aliments

Phytic acid is naturally present in feedstuffs of plant
origin as phytate, where it serves as the storage form of
phosphorus (Bedford 2000). However, phytic acid is
negatively charged at acidic, neutral and basic pH
conditions (Maenz 2001), and hence it can bind posi-
tively charged molecules in the diet and in endogenous
gastrointestinal tract secretions such as digestive en-
zymes and mucins at all pH conditions found in the
gastrointestinal tract, thereby reducing nutrient digesti-
bility and increasing the endogenous secretion of
nutrients. Phytase enzyme, which hydrolyses phytic
1 Adeola and Sands 2003; Selle et al. 2006; Cowieson
Corresponding author. Current address: Department of
Agricultural, Food and Nutritional Science, University of
Alberta, Edmonton, Alberta, Canada T6G 2P5.
acid, may reduce the negative effects of phytic acid.
The addition of non-starch polysaccharides degrading
carbohydrases to phytase-supplemented diets may en-
hance the efficacy of phytase by reducing the negative
effects of phytic acid because the carbohydrases may
hydrolyze the non-starch polysaccharides to increase the
accessibility of phytase to phytic acid (Woyengo and
Nyachoti 2011).
The effect of dietary phytic acid and supplemental
phytase on nutrient digestibility has been investigated in
several studies and it has been reviewed (Thompson
1986; Pallauf and Rimbach 1997; Sebastian et al. 1998;
Abbreviations: ENL, endogenous nutrient losses; PA, phytic acid

Can. J. Anim. Sci. (2013) 93: 921 doi:10.4141/CJAS2012-017 9

 10 CANADIAN JOURNAL OF ANIMAL SCIENCE

Higherperformance of animals fed the control diet was higher than that of animals fed the phytic acid diet; Similar performance of animals fed the control was similar to that of animals
Onyango and Adeola (2009)
et al. 2011). Also, the effect of adding non-starch
polysaccharides degrading carbohydrases to phytase-

Shan and Davis (1994)

Woyengo et al. (2012)

Cabahug et al. (1999)
supplemented diets on nutrient digestibility has been

Ceylan et al. (2003)

Reference
determined in several studies, and we have reviewed data

Liu et al. (2008b)

Liu et al. (2008a)
Liu et al. (2009)
from these studies (Woyengo and Nyachoti 2011).
However, information on the effect of phytic acid on
the endogenous losses of nutrients in pigs and poultry,
and on the mechanisms by which phytic acid increases
the endogenous nutrient losses has not been reviewed.
Therefore, there is a need to review this information

Decrease in performance
and integrate it with that on the effect of phytic acid on

due to PA (%)
nutrient digestibility in order to gain a broader under-
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standing of the mechanisms by which phytic acid reduces

28
44

37
3
3
7
7


nutrient utilization in pigs and poultry as well as the
effectiveness of possible interventions. The objective of
this paper is, therefore, to review results from studies on
the effects of phytic acid on performance and nutrient
utilization of pigs and poultry, and to discuss mechan-

control vs. PA dietx

isms by which phytic acid reduces nutrient utilization in

Performance of
these animals. Also, areas that need further research

Similar
Higher
Higher
Higher
Higher
Higher
Higher

Higher
in order to gain more insight into mechanisms by which
phytic acid reduces nutrient utilization in pigs and
poultry are suggested. The chemical composition and
the occurrence of phytic acid in feedstuffs of plant origin
have been discussed in detail in our previous review
paper (Woyengo and Nyachoti 2011), and hence they are

Egg production
For personal use only.

not covered in the current review.

Response
criteriony
Table 1. Results from studies on the effect of dietary phytic acid on performance of poultry and pigs

EFFECT OF PHYTIC ACID ON ANIMAL

BWG
BWG
BWG
BWG
BWG
BWG

BWG
PERFORMANCE
The presence of phytic acid in diets for poultry and pigs
PA content in

has generally been shown to result in reduced growth

PA diet (%)

performance (Table 1). Thus far, only one study has

1.57
1.57
1.57
1.57
1.65
1.65
0.71
been conducted to investigate the effect of dietary phytic 2.00
acid on egg production by laying hens (Table 1). In this
study, there was no effect of increasing dietary phytic
control diet (%)

acid from 0.57 to 0.71% on egg production, which may

Study duration PAz content in

have been due to the fact that the increase in dietary

0.78
0.78
0.78
1.04
0.00
0.00
0.57
0.00

phytic acid level from 0.57 to 0.71% was not high

enough to impact egg production. In the broiler and pig
studies in which an increase in dietary phytic acid
resulted in reduced performance, the dietary phytic
acid level had been increased by more than 0.5
percentage points (Table 1). Therefore, there is a need
21
28
21
18
14
18

21
140
(d)

for research on the effect of dietary phytic acid on

performance of laying hens fed diets in which dietary
phytic acid level has been increased by more than 0.5
percentage points. Also, there is a need for research on
the effect of dietary phytic acid on turkey performance.
Initial age (d)

Supplemental phytase has been added to pig and

Chicks of a laying strain.
BWG, body weight gain.
0
0
0
7
8

140
28

25

poultry feeds to improve phosphorus availability by

fed the phytic acid diet.

hydrolyzing phytic acid (Selle and Ravindran 2007,

2008). However, reduced performance of pigs and
PA, phytic acid.

poultry due to phytic acid implies phytase may be added

Animal type

Laying hens

to diets to improve performance of swine and poultry

not only by improving phosphorus availability, but by
Chicksw
Broiler
Broiler
Broiler
Broiler
Broiler

Piglets

alleviation of the other anti-nutritional effects of phytic

acid as well (Woyengo et al. 2012).
w
x
y
z
 WOYENGO AND NYACHOTI * ANTI-NUTRITIONAL EFFECTS OF DIETARY PHYTIC ACID 11
MECHANISM OF ACTION OF PHYTIC ACID
AGAINST GROWTH PERFORMANCE
Effect of Phytic Acid on Apparent Mineral
Digestibility
The effect of phytic acid on apparent mineral digest-
ibility in pigs and broilers has been discussed in previous
review papers (Cowieson et al. 2011; Woyengo and
Nyachoti 2011). In summary, phytic acid has been
shown to reduce the apparent mineral digestibility in
pigs and broilers. However, Ceylan et al. (2003) reported
that an increase in dietary phytic acid from 0.57 to
0.71% did not reduce the retention of cationic minerals
Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.230.235 on 05/17/20
by laying hens fed diets in which non-phytate phospho-
rus level ranged from 0.2 to 0.4%. They instead
observed an interaction between dietary phytic acid
and non-phytate phosphorus such that the retention of
the cationic minerals increased with an increase in
dietary non-phytate phosphorus level when dietary
phytic acid level was 0.71%, and decreased with an
increase in dietary non-phytate phosphorus level when
dietary phytic acid level was 0.57%. Thus, the effect of
dietary phytic acid on the retention of the cationic
minerals by the laying hens in this study of Ceylan et al.
(2003) was confounded by dietary non-phytate phos-
phorus. Also, like performance, the lack of effect of
phytic acid on the cationic mineral retention could have
For personal use only.
been due to the fact that the increase in dietary phytic
acid level from 0.57 to 0.71% was not high enough to
impact mineral retention by the laying hens. Hence,
there is a need for more research on the effect of phytic
acid on mineral digestibility by laying hens fed diets
in which dietary phytic acid has been increased by a
magnitude that is greater than that by which it was
increased in the study of Ceylan et al. (2003). Also, there
is lack of information on the effect of dietary phytic acid
on mineral digestibility in turkeys, and hence there is a
need to fill this gap in knowledge.
The reduced apparent mineral digestibility by phytic
acid could be due to reduced availability of dietary
minerals for absorption or increased endogenous secre-
tion of the minerals in the gastrointestinal tract lumen or
both. The effects of phytic acid on mineral absorption
and secretions are discussed below.
Effect of Phytic Acid on True Absorption of
Minerals
The effect of phytic acid on the absorption of minerals
has been investigated in humans and rats. In humans,
Bronner et al. (1954) reported a 55% decrease in 45Ca
absorption in adolescent boys fed farina meal due to an
increase in phytic acid concentration in their diet from 0
to 0.5%. Similarly, Kim et al. (2007) reported reduced
zinc absorption in young women (43 vs. 22%) and in
elderly women (34 vs. 20%) due to an increase in dietary
phytate intake from 690 to 1623 mg d 1 and from 760
to 1713 mg d1, respectively. Hunt and Beiseigel (2009)
also reported a 25% or 1 mg d 1 decrease in absolute
zinc absorption in women due to an increase in
their dietary intake of phytic acid from 440 and
1800 mg d1.
In rats, Lönnerdal et al. (1989) observed an increase in
the amount of dietary 45Ca in the cecal contents of the
rats from 0.45 to 17.5% due to dietary phytic acid,
indicating reduced absorption of calcium in rats due to
phytic acid. Kim et al. (1993) also reported a 33%
decrease in dietary 59Fe3absorption in rats fed a cereal
meal due to addition of 0.13% phytic acid to the meal.
With regards to zinc absorption, Rubio et al. (1994)
reported an 18% reduction in absorption of 65Zn in rats
fed a phytic acid-free diet after supplementing the diet
with phytic acid at 0.3%. Similarly, Lönnerdal et al.
(1989) reported an 84% reduction in dietary 65Zn
uptake by rats’ liver due to including phytic acid in
the solution fed, indicating that phytic acid reduced
absorption of zinc in rats. In an in situ study with rats,
Davies and Nightingale (1975) reported a decrease in
65
Zn absorption in ligated loops of rat duodenum (con-
taining 1 mL of a saline solution with 5 mg of 65Zn) by
96% due to the addition of 5.07 mg of phytic acid into
the solution. There is, however, a lack of information on
the effect of phytic acid on true absorption of minerals
in pigs and poultry.
In summary, phytic acid can reduce the absorption of
dietary minerals leading to reduced mineral availability
to the animals, which could partly explain the reduced
animal performance due to phytic acid. The mechanism
by which phytic acid can reduce the availability of
minerals for absorption has been suggested (Woyengo
and Nyachoti 2011). Phytic acid is poorly hydrolyzed by
pigs and poultry and it can form insoluble complexes
with cations at neutral pH found in the small intestine
(Prattley et al. 1982; Woyengo and Nyachoti 2011). For
example, Lyon (1984) reported that the addition of
phytic acid to a mineral solution at neutral pH resulted
in precipitation of 99.5, 75, 83 and 62% of zinc, iron,
calcium and magnesium in the solution, respectively,
whereas Davies and Olpin (1979) reported that the
addition of phytic acid to a mineral solution at neutral
pH resulted in precipitation of 98, 91, and 80% of zinc,
copper and manganese in the solution, respectively.
However, sodium forms weaker bonds with phytic acid
than divalent cations (Erdman 1979), and hence its
solubility is not affected by phytic acid (Scheuermann
et al. 1988). Therefore, it appears that the availability
of monovalent cations such as sodium for absorption
is not affected by phytic acid. However, there is lack of
information on the effect of phytic acid on true
absorption of monovalent cations in the gastrointestinal
tract of non-ruminants.
Effect of Phytic Acid on Endogenous Losses of
Minerals
The effect of phytic acid on endogenous losses of
minerals has been investigated. For example, Woyengo
et al. (2009) observed a decrease in the apparent ileal
 12 CANADIAN JOURNAL OF ANIMAL SCIENCE
digestibility of sodium and magnesium in piglets to
negative values (18.2 and 3.0%, respectively) as a
result of increasing the dietary content of phytic acid (as
sodium phytate) from 0 to 2%, indicating increased
endogenous flow of this mineral at the terminal ileum of
pigs due to phytic acid. In rats, phytic acid was shown to
reduce the biological half-life for body 65Zn, indicating
that phytic acid increased the endogenous losses of zinc
(Davies and Nightingale 1975).
In poultry, Cowieson et al. (2004) observed a 68, 32,
300 and 47% increase in endogenous excretion of
calcium, iron, sodium and sulfur, respectively, in
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broilers that had been ‘‘precision fed’’ 50 mL of a
glucose solution over a 48-h period due to addition of
1 g of phytic acid to the 50 mL of glucose solution.
Cowieson et al. (2006) ‘‘precision fed’’ broilers a
solution containing 5 g of casein with or without 1 g
of phytic acid and also reported increased excretion of
calcium, magnesium, manganese, and sodium by 187,
39, 87, and 174%, respectively, over a 48-h period, due
to phytic acid. However, Liu and Ru (2010) did not
observe any effect of increasing the dietary content of
phytic acid from 0.71 to 1.42% on the ileal endogenous
flows of manganese, zinc, magnesium, potassium and
calcium in broilers. Instead, they reported 15 and 18%
decreases in ileal endogenous flows of iron and copper,
For personal use only.
respectively, and the reason why results from their study
were different from those studies of Cowieson et al.
(2004, 2006) is not clear. It should, however, be noted
that the phytic acid used in the study of Liu and Ru
(2010) was in the form of sodium salt (sodium phytate),
whereas that used in the studies of Cowieson et al. (2004,
2006) was a free phytic acid. Onyango et al. (2009)
determined the effect of form of phytic acid (free phytic
acid vs. magnesiumpotassium phytate) on endogenous
losses of mucin and amino acids in broilers and observed
that the extent to which phytic acid increased the
endogenous losses of the same was dependent on the
form of phytic acid; magnesium-potassium phytate was
more potent. Therefore, the differences between the
study of Liu and Ru (2010) and those of Cowieson et al.
(2004, 2006) with regard to the effects of phytic acid on
endogenous losses of minerals might have been due to
the differences between the forms of phytic acid used in
the studies.
Although the dietary form of phytic acid (free phytic
acid vs. phytate salt) can influence the effect of phytic
acid on nutrient utilization, this influence of the form of
phytic acid on nutrient utilization may occur only in
poultry (and not in pigs) because of the following three
reasons. First, phytic acid and the cationic minerals
disassociate at the acidic pH found in the stomach
because they are both soluble at this pH (Maenz 2001).
Therefore, the form in which phytic acid exerts its effects
in the stomach is independent of the original form
(free phytic acid vs. phytate salt) in which it is sup-
plied in the diet. Second, at small intestinal pH, phytic
acid reacts with free cationic minerals to form phytate
(Maenz 2001). The major cationic mineral in practical
diets is calcium. Therefore, phytic acid is likely to
complex more calcium than other cations regardless of
its original form in the diet. Third, the pH in the crop of
poultry is not acidic enough to cause disassociation
between phytic acid and cationic minerals, and therefore,
phytic acid may exert its effects in the crop in a form that
is similar to the form that it is supplied in the diet,
resulting in an overall influence of dietary form of phytic
acid on response to its ingestion. However, there is a need
for research on the effect of form of phytic acid on
nutrient utilization in poultry and pigs with the view of
testing this hypothesis.
From these studies on the effect of phytic acid on
endogenous losses of minerals, it is apparent that phytic
acid increases the endogenous losses of minerals. Four
mechanisms by which phytic acid can increase the
endogenous secretion of minerals are proposed. First,
phytic acid may interact with dietary protein and
endogenous protein (pepsin, and pepsinogen and its
activating peptide) in the stomach, where the pH is
acidic (Cowieson et al. 2006), thereby reducing the
activity of pepsin. Indeed, Knuckles et al. (1989)
reported a 9 to 14% reduction in pepsin digestion of
casein and bovine serum albumin in vitro due to
addition of phytic acid to the incubation media, whereas
Vaintraub and Bulmaga (1991) reported a 60 to 92%
reduction in pepsin digestion of casein, hemoglobin,
bovine serum albumin and soybean protein in vitro due
to addition of phytic acid to the incubation media.
Furthermore, Liu et al. (2009) reported a 6.3% reduc-
tion in the activity of pepsin in the proventriculus of
broiler chickens due to dietary phytic acid, whereas
Woyengo et al. (2010) observed reduced activity of
pepsin in piglets by 46% due to addition of phytic acid
to a phytic acid-free diet.
The presence of undigested feed or inactivation of
digestive enzymes in the gastrointestinal tract results in
increased secretion of the latter through negative feed-
back mechanisms (Hara et al. 2000; Morisset 2008).
Binding of tannic acid to pepsin in the stomach of rats
was shown to result in increased pepsin and hydrochlo-
ric acid secretions (Mitjavila et al. 1973). Therefore, the
binding of phytic acid to pepsinogen and its activating
peptide in the stomach can result in an increase in the
secretion of the pepsinogen and hydrochloric acid via
negative feedback mechanisms. Neutralization of the
resulting acidic digesta by intestinal (Allen et al. 1993)
and pancreatic (Zebrowska et al. 1983) bicarbonates for
small intestinal mucosa protection against acidity and
digestive enzyme activity optimization increases the loss
of endogenous minerals. Pancreatic juice is richer in
sodium than other minerals (Zebrowska et al. 1983).
Therefore, it appears that sodium is the mineral whose
endogenous secretion is most dramatically affected by
increased secretion of pepsinogen and hydrochloric acid
due to phytic acid.
 WOYENGO AND NYACHOTI * ANTI-NUTRITIONAL EFFECTS OF DIETARY PHYTIC ACID 13
Second, several digestive enzymes in the gastrointest-
inal tract including a-amylase, alkaline phosphatase,
aminopeptidases and carboxypeptidases are metalloen-
zymes (Malmstrom and Neilands 1964; Wouters and
Husain 2001). For instance a-amylase is a metalloen-
zyme that requires calcium as a co-factor (Argent et al.
1973), whereas alkaline phosphatase, aminopeptidases
and carboxypeptidases are metalloenzymes that require
zinc as a co-factor (Malmstrom and Neilands 1964;
Wouters and Husain 2001). These metal ions are tightly
bound to the enzymes to the extent that they are an
integral part of the structure of the enzyme (Malmstrom
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and Neilands 1964), meaning that an increase in
secretion of a metalloenzyme will automatically lead to
increased secretion of its co-factor. An increase in
secretion of pancreatic a-amylase in the pancreatic juice
has indeed been shown to result in an increased secretion
of calcium (Argent et al. 1973). Therefore, phytic acid
can bind dietary protein or endogenous protein (diges-
tive enzymes) or both through multivalent cations in the
small intestine to form insoluble complexes (Prattley
et al. 1982; Thompson 1986; Maenz 2001), leading to
reduced activity and hence increased secretion of the
digestive enzymes. The ability of phytic acid to reduce
the activity of digestive enzymes found in the small
intestine was demonstrated by Deshpande and Cheryan
For personal use only.
(1984), who observed reduced a-amylase activity in vitro
due to phytic acid, and by Liu et al. (2008b), who
reported an 8.3% reduction in a-amylase activity in the
duodenum of broilers due to dietary phytic acid.
Therefore, phytic acid may bind to enzyme co-factors,
enzymatic protein, dietary protein, or all of them,
thereby increasing the secretion of the co-factors
through negative feedback mechanisms. Of all the
endogenously secreted minerals, calcium and zinc
are most affected by this process; this is because they
are the co-factors of metalloenzymes found in the
gastrointestinal tract (Malmstrom and Neilands 1964).
Third, phytic acid can bind to endogenously secreted
minerals thereby preventing their re-absorption. Multi-
valent cations form insoluble complexes with phytic acid
(Maenz 2001), meaning that phytic acid can reduce
their re-absorption. However, monovalent cations form
weaker bonds with phytic acid than their multivalent
counterparts (Erdman 1979; Scheuermann et al. 1988),
and hence the re-absorption of the endogenously
secreted monovalent cations is less likely to be affected
by binding to phytic acid than multivalent cations.
Fourth, the active transport of nutrients (solutes) into
enterocytes generates an osmotic flow of water into the
enterocytes which, in turn, results in an increase in
absorption of minerals by solvent drag (Fordtran et al.
1968). Sodium is additionally absorbed from the small
intestine by co-transportation with other nutrients
including glucose and galactose (Fordtran et al. 1968),
and its absorption was shown to increase with an
increase in glucose absorption (Fordtran 1975; Schiller
et al. 1997). Phytic acid has been shown to reduce true
nutrient digestibility or absorption. Therefore, phytic
acid may reduce the re-absorption of all the endogen-
ously secreted minerals by reducing the absorption
of nutrients such as sugars and amino acids in the
gastrointestinal tract.
In summary, the increased endogenous loss of miner-
als due to phytic acid could be a result of their increased
secretion in the small intestine or their reduced re-
absorption from the small intestine or both (Fig. 1).
It appears that the increased endogenous loss of sodium
due to phytic acid is mainly due to increased secretion as
a result of reduced pepsin activity and hence increased
secretion of pepsin and hydrochloric acid in the stomach
and the consequent increase of mineral bicarbonates to
neutralize the acid; or reduced re-absorption due to
reduced absorption of other nutrients, whose absorption
results in increased absorption of sodium; or both.
However, there is a need to test these hypotheses.
The increased endogenous losses of minerals due to
phytic acid may partly explain the reduced performance
of animals fed phytic acid-supplemented diets. This is
because an increase in endogenous losses of nutrients
can result in increased dietary demand of the lost
nutrients by the animal and of the energy required to
secrete them (Nyachoti et al. 1997). Also, sodium
deficiency in chickens has been reported to reduce the
activity of NaKATPase in the gastrointestinal tract
and to increase the expression of genes for the same
enzyme which is involved in the absorption of several
nutrients including glucose and amino acids (Gal-
Garber et al. 2003). Therefore, the increased secretion
of sodium may alter the synthesis and activity of
ATPases, which are involved in nutrient absorption.
Liu et al. (2008b) have shown reduced activity of NaK
ATPase in broilers due to ingestion of phytic acid. We
have also observed reduced activity of NaKATPase in
the jejunum of piglets by 41% due to ingestion of phytic
acid (Woyengo 2010).
Effect of Phytic Acid on Apparent Nitrogen and
Amino Acid Digestibilities
The apparent digestibility of nitrogen and amino acids
in broilers and pigs has been reported to be reduced due
to an increase in dietary concentration of phytic acid
(Table 2). However, Knuckles et al. (1989) did not
observe any effect of increasing the level of phytic acid
in casein-cornstarch-based diets for rats from 0 to 2%
on apparent ileal protein digestibility. We either did not
observe any effect of supplementing a casein-cornstarch-
based (phytic acid-free) diet with phytic acid at 2% on
apparent ileal amino acids in weanling pigs (Woyengo
et al. 2009).
Pigs and poultry differ in digesta dry matter content
and rate of passage of digesta in their small intestines,
and these differences have been suggested to be the
causes of differences between pigs and poultry with
regard to their susceptibility to anti-nutritional factors
(Bedford and Schulze 1998). Therefore, differences in
 14 CANADIAN JOURNAL OF ANIMAL SCIENCE

Dietary phytic acid

Binding of dietary phytic acid to Binding of dietary phytic acid to

nutrients and digestive enzymes dietary protein and digestive
in small intestine enzymes in stomach

Binding of dietary phytic acid

Reduced activity of digestive to endogenous nutrients Reduced pepsin
enzymes in small intestine activity in stomach
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Increased secretion of
pepsin and HCl in stomach

Increased acidity of
digesta flowing to duodenum

Increased secretion of mineral

Increased secretion of digestive
enzymes in small intestine
For personal use only.
Reduced re-absorption of
bicarbonates and mucins in
endogenous nutrients
small intestine

Increased ileal endogenous

flow of nutrients

Reduced true ileal

nutrient digestibility

Fig. 1. Proposed mechanism of action of dietary phytic acid on ileal digestibility and endogenous losses of nutrients.

the composition of basal diets may also cause differences
in pigs or poultry with regard to the susceptibility to
anti-nutritional factors because diet composition is
known to affect viscosity and rate of passage of digesta
in the gastrointestinal tract of pigs and poultry (Bedford
2000). The basal diets used in the our study (Woyengo
et al. 2009) and that of Knuckles et al. (1989) in which
dietary phytic acid did not affect amino acid digestibility
in pigs were based on cornstarch and casein, whereas
diets used in the studies of Bohlke et al. (2005) and Liao
et al. (2005) in which dietary phytic acid reduced amino
acid digestibility in pigs (see Table 2) were based on corn
and soybean meal. Therefore, the differences in the
effect of dietary phytic acid on amino acid digestibilities
among the studies may have been due to differences in
the composition of basal diets. However, there is a need
to confirm that cornstarchcasein-based and corn
soybean meal-based diets have different effects on
digesta dry matter content and rate of passage of digesta
in small intestine of pigs or poultry. Also, there is a need
to establish the mechanisms by which digesta dry matter
content and rate of passage of digesta in the small
intestine could affect the influence of dietary phytic acid
on amino acid digestibility by pigs or poultry.
There is lack of information on the effect of dietary
phytic acid on amino acid digestibility in laying hens
and turkeys. Hence, there is a need to fill this gap in
knowledge.
Like for minerals, the reduced apparent amino acid
digestibility due to phytic acid could be due to reduced
availability of dietary amino acids for absorption or
increased endogenous losses of amino acids or both.
These factors are discussed below.
Effect of Phytic Acid on True Digestibility and
Absorption of Amino Acids
Phytic acid supplementation was shown to decrease
true ileal digestibility of amino acids by at least 8%
(Cowieson et al. 2006) and true metabolizable nitrogen
by at least 120% (Cowieson et al. 2003) due to the
 WOYENGO AND NYACHOTI * ANTI-NUTRITIONAL EFFECTS OF DIETARY PHYTIC ACID 15

addition of 0.5 g of phytic acid to 5 g of casein that had

Change
been precision fed to growing broilers. Onyango et al.

NS
NS

NS
S
S

S
S
S
S

Liao et al. (2005) (2008) perfused the jejunum of broilers with a solution
containing labeled amino acids to determine the effect of
phytic acid on absorption of amino acids. They reported
1.56
a linear decrease in absorption of a labeled leucine due

77.8
85.2
80.8
75.2

76.8

77.3
64.7
71.4
72.1
NR
Diet PA (%)

to an increase in magnesium-potassium phytate concen-

tration in the perfusate from 0 to 500 mM. In their
study, the phytate also tended to linearly reduce the
79.9
0.78

85.9
84.8
78.9

78.2

81.5
69.3
76.0
75.2
absorption of lysine and glutamic acid. We, however,

NR
Growing pig

did not observe any effect of supplementing a casein-

cornstarch-based (phytic acid-free) diet with phytic acid
Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.230.235 on 05/17/20

at 2% on true ileal amino acid digestibilities in weanling

Change

pigs (Woyengo et al. 2009), which may have due to

NSv

NS

NS

NS
S

S
S
S
differences between species (pigs vs. poultry) with regard
Bohlke et al. (2005)

to digesta dry matter content and rate of passage of

digesta in small intestine as previously discussed.
Phytic acid may exert its detrimental effects on true
76.6

83.3
75.1

67.9

57.4
79.5
77.8
57.0
61.6
64.5
NR
Diet PA (%)

amino acid digestibility or absorption by binding to

Table 2. Apparent digestibility of indispensable amino acids and nitrogen by broilers and growing pigs due to dietary phytic acid

dietary protein and proteolytic enzyme proteins in the

stomach and in the small intestine, thereby reducing the
x

availability of protein for digestion in the stomach and

78.4

84.5
79.6

71.2

65.5
79.7
80.8
61.8
67.4
68.2
NR

the small intestine (Reddy et al. 1989; Fig. 1). In

addition to reducing pepsin activity, Liu et al. (2009)
have reported a 24.2% reduction in trypsin activity in
the jejunum of chickens due to an increase in dietary
Change
For personal use only.

NS

NS

phytic acid level from 0.78 to 1.56%. Thus, phytic acid

S

S
S

S
S
S
Ravindran et al. (2006)

can reduce performance partly by reducing the digest-

ibility of dietary amino acids.

Effect of Phytic Acid on Endogenous Losses of

74.2

81.2
1.36

86.6

79.5

84.5
89.4
80.4
73.3
76.9
80.7
Diet PA (%)

Amino Acids
Change change in digestibility due to an increase in dietary phytic acid content.

Studies on the effect of phytic acid on endogenous losses

of nitrogen and amino acids by broilers generally show
79.0

83.4
1.04

88.9

82.0

87.3
89.6
83.2
77.3
80.5
81.7

an increase in endogenous losses of amino acids with

increasing dietary phytic acid (Table 3). However, there
Broiler

is a lack of information on the effect of dietary phytic

acid on the effect of dietary phytic acid on endogenous
y

amino acid losses in laying hens and turkeys. Hence,

Change
w

S
S

S
S
S
S


there is a need to fill this gap in knowledge.

S
Ravindran et al. (2000)

In pigs, we did not observe any effect of supplement-

ing a caseincornstarch-based (phytic acid-free) diet
with phytic acid at 2% on ileal endogenous losses of
amino acids in weanling pigs (Woyengo et al. 2009).
80.5

76.9
1.57

85.8

79.4

84.4

79.6
72.1
78.0
80.7
NR
Diet PAz (%)

This indicates that phytic acid has variable effects not

only on amino acids digestibility in pigs but also on
Diet PA, dietary phytic acid content.

endogenous loss of amino acids in the same species.

82.1

78.5
1.04

86.5

81.3

85.8

81.5
74.6
79.8
83.0
NR

However, so far, there is no other study that has

reported the effect of phytic acid on endogenous loss
of amino acids in pigs. Therefore, it is difficult to draw a
firm conclusion on the effect of phytic acid on endo-
genous loss of amino acids in pigs. More research needs
NS, non-significant.
NR, not reported.

to be done on the effect of phytic acid on endogenous

loss of amino acids in pigs.
S, significant.
Phenylalanine

We have observed reduced gastric pepsin activity in

Methionine

Threonine
Isoleucine
Item (%)

Histidine

Nitrogen
Arginine

piglets due to phytic acid (Woyengo et al. 2010),

Leucine
Lysine

Valine

implying that phytic acid increases pepsin secretion in

pigs. Therefore, the lack of effect of phytic acid on the
w
x
y
z

v
 16
CANADIAN JOURNAL OF ANIMAL SCIENCE
Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.230.235 on 05/17/20

Table 3. Increase in endogenous losses of indispensable amino acids and nitrogen by broilers due to dietary phytic acid

Liu and Ru Cowieson et al. Cowieson and Ravindran Cowieson et al. Onyango et al. Onyango et al.
Item (2010) (2008) (2007) (2004) (2009) (2009)

Animal age 21 28 26 42 70 70
PAz content in control diet (%) 0.71 0.85 0.00 0.00 0.00 0.00
PA content in PA diet (%) 1.42 1.45 1.45 2.00 2.00 2.00
Supplemental PA form Na-phytate Na-phytate Na-phytate Free PA Free PA Mg-K-phytate
Method of feedingy Ad-libitum Ad-libitum Ad-libitum Precision Precision Precision
For personal use only.

Unit of reduction mg kg 1 DMx mg kg1 DM mg kg1 DM mg bird 1 48 h 1 mg bird 1 54 h 1 mg bird 1 54 h 1

Arginine 83 123 228 17 12 11
Histidine 1 203 73 15 9 10
Isoleucine 11 142 435 22 5 6
Leucine 80 386 350 35 10 9
Lysine 23 120 219 16 12 8
Methionine 40 37 89 0 2 3
Phenylalanine 21 279 71 21 6 6
Threonine 27w 349 866 19 16 15
Valine 121 143 324 28 8 7
Nitrogen Not reported 980 1251 Not reported Not reported 11

z
PA phytic acid.
y
In the study of Cowieson et al. (2004), the birds were ‘‘precision fed’’ 50 mL glucose solution, whereas in the study of Onyango et al. (2009), the birds were ‘‘precision fed’’ 50 mL dextrose
solution.
x
DM, dry matter.
w
Negative value indicate that dietary phytic acid reduced the endogenous loss of threonine in the study of Liu and Ru (2010).
 WOYENGO AND NYACHOTI * ANTI-NUTRITIONAL EFFECTS OF DIETARY PHYTIC ACID 17
endogenous loss of amino acids in pigs may be attri-
buted to the fact that phytic acid increases the endo-
genous secretion of amino acids, but it does not affect
their absorption, leading to efficient re-absorption of the
endogenously secreted amino acids. However, it should
be noted that energy is spent during the synthesis,
secretion and re-absorption of endogenous protein. This
implies that the reduced pepsin activity by phytic acid
may result in increased energy expenditure in gastro-
intestinal and hence reduced pig performance without
an effect on amino acid digestibility. Therefore, there is
a need to determine the effect of phytic acid on energy
Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.230.235 on 05/17/20
expenditure in the gut relative to the whole body in
pigs to establish whether dietary phytic acid can increase
energy expenditure in the gastrointestinal tract. There is
also a need to determine the effect of phytic acid on
energy expenditure in the gut relative to the whole
body in poultry to establish whether the increased
endogenous loss of amino acids due to dietary phytic
acid can result in increased energy expenditure in gut.
Endogenous nitrogen and amino acids in the gastro-
intestinal tract originate from digestive enzymes, muco-
proteins and sloughed cells of gastrointestinal tract
epithelium (Nyachoti et al. 1997). The amounts of
endogenous amino acids that appear at the terminal
ileum are a proportion of what is secreted, but not re-
For personal use only.
absorbed (Nyachoti et al. 1997). Thus, the increased
endogenous flow of amino acids at the terminal ileum
might be a result of their increased secretion or reduced
re-absorption or both. Phytic acid may increase the
endogenous secretion of amino acids by binding to
dietary protein and endogenous digestive enzymes in the
gastrointestinal tract, thereby increasing the secretion of
the enzymes through negative feedback mechanisms.
The increased secretion of digestive enzymes may in
turn result in increased secretion of mucins, leading to
further increase in endogenous losses of amino acids.
This is because mucins are partly composed of protein
(Forstner and Forstner 1994), and an increase in the
secretion of gastric pepsin has been shown to result in
increased secretion of mucins to protect the gastro-
intestinal tract mucosa from digestion by the enzyme
and by hydrochloric acid, whose secretions increase
with increases in pepsin secretion (Munster et al. 1987).
Cowieson et al. (2004) have observed increased secretion
of sialic acid, which is a major component of mucin, in
broilers due to dietary phytic acid. Onyango et al. (2009)
have also observed increased secretion of sialic and
mucins in broilers due to dietary phytic acid, indicating
that the secretion of mucins is indeed increased by phytic
acid.
In addition to increasing the secretion of digestive
enzymes and mucin, phytic acid may increase the
endogenous losses of amino acids by reducing the
re-absorption of the endogenously secreted amino acids
in the small intestine. This is because phytic acid has
been shown to reduce true amino acids digestibility or
absorption in poultry, implying that it can also reduce
the digestibility of the endogenously secreted proteins
and hence the re-absorption of the endogenous amino
acids.
Therefore, it appears that phytic acid increases the
endogenous amino acid losses by increasing the secre-
tion of digestive enzymes and mucins, and by reducing
the re-absorption of the endogenously secreted amino
acids in the small intestine (Fig. 1). Because the
increased endogenous nitrogen and amino acid losses
in the gastrointestinal tract are associated with increased
maintenance requirement of the lost nitrogen and amino
acids and of the energy spent on their secretion
(Nyachoti et al. 1997), an increase in endogenous losses
of nitrogen and amino acids due to phytic acid imply
that the presence of the phytic acid in diets for non-
ruminants results in increased maintenance require-
ments of energy and amino acids, thereby reducing the
availability of these nutrients for tissue deposition.
Therefore, the reduced animal performance due to
dietary phytic acid could partly be explained by the
increased endogenous losses of nitrogen and amino
acids.
Effect of Phytic Acid on Energy Digestibility
Liao et al. (2005) reported reduced apparent ileal and
total tract digestibilities of energy in weanling pigs by
7.5 and 6.1%, respectively, due to an increase in dietary
phytic acid concentration from 0.78 to 1.70%. In
poultry, Ravindran et al. (2006) reported decreased
apparent metabolizable energy content in broiler diets
by 2.0% due to an increase in dietary concentration of
phytic acid by 36%.
Phytic acid may decrease energy digestibility by
reducing the digestibility of energy generating molecules
such as carbohydrates, lipids and protein. For example,
Thompson et al. (1987) reported that dephytinization
of navy beans resulted in a 25% increase in starch
digestibility in vitro. They also observed decreased car-
bohydrate malabsorption (estimated by breath hydro-
gen) and improved glycemic index in human subjects
by 81 and 126%, respectively, due to consumption
of bread made from dephytinized navy beans. In mice,
Lee et al. (2006) reported reduced fasting and non-
fasting blood glucose levels in male diabetic mice fed a
casein-corn starch-based diet by 21 and 15%, respec-
tively, due to an increase in the dietary level of
phytic acid from 0 to 1.0%, indicating reduced glucose
absorption as a result of the dietary phytic acid.
In broilers, an increase in dietary phytic acid level
from 0.79 to 1.57% was shown to result in a decrease
in blood serum glucose concentration by 6% (Liu et al.
2008b), indicating reduced glucose absorption due to the
phytic acid. Onyango et al. (2008) also observed a
tendency for phytic acid to linearly reduce the absorp-
tion of glucose in jejunum of broilers after perfusion of
the jejunum with a solution that contained a labeled
glucose and phytic acid at increasing concentrations
(0, 50, 250 or 500 mM).
 18 CANADIAN JOURNAL OF ANIMAL SCIENCE
Deshpande and Cheryan (1984) observed reduced
activity of a-amylase in vitro due to phytic acid, whereas
Liu et al. (2008b) reported reduced activities of a-
amylase, sucrase and maltase in the duodenum of
broilers by 8.3, 11.4 and 6.0%, respectively. Thus, it
appears that phytic acid reduces the absorption of
carbohydrates likely by reducing the activity of digestive
carbohydrases. Phytic acid may reduce activity of
digestive carbohydrases by binding to: (i) the digestive
enzymes, (ii) dietary protein that is closely associated
with starch, and (iii) starch through phosphate linkages
(Thompson 1986). The reduced glucose absorption by
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phytic acid could partly explain the reduced energy
digestibility by phytic acid. However, research is re-
quired to probe the effect of phytic acid on digestibility
or absorption of carbohydrates in pigs.
In studies on lipid digestibility, Lee et al. (1997)
reported increased fecal cholesterol and bile acid
excretion in rats due to dietary phytic acid, indicating
that phytic acid reduced the (re)absorption of choles-
terol and bile acids. Lee et al. (2005) fed diabetic mice
diets containing 0, 1 and 1.5% phytic acid to investi-
gate the effect of phytic acid on the levels of lipids in
the serum and liver. They observed lower serum total
cholesterol and low-density lipoprotein cholesterol
concentrations as well as hepatic total lipid and total
For personal use only.
cholesterol due to an increase in the level of dietary
phytic acid from 0 to 1.5%. Similarly, Lee et al. (2007)
reported lower serum low-density lipoprotein choles-
terol and hepatic triacylglycerol in aged mice due to an
increase in dietary phytic acid from 0 to 1.5%. In their
study, the apparent absorption of total lipid and
cholesterol was also reduced, whereas fecal cholesterol
and lipid contents were increased by the dietary phytic
acid. Yuangklang et al. (2005) also reported increased
excretion of bile acids in feces and reduced concentra-
tion of cholesterol in the liver of rats fed a cholesterol
rich diet by 28 and 22%, respectively, due to an
increase in dietary phytic acid concentration from
0 to 0.33%. However, the increased fecal excretion of
bile acids did not translate to reduced fat digestibility
and serum cholesterol concentration and the reason for
this is not clear. It could probably be attributed to the
fact that the dietary level of phytic acid (0.33%) was
not adequate or enough to impact fat digestibility.
In poultry, Liu et al. (2009) reported reduced serum
total cholesterol and hepatic triacylglycerol content due
to an increase in the level of phytic acid level in the diets
for broilers from 0.7 to 1.4%. Thus, results from these
studies indicate that phytic acid reduces the (re)absorp-
tion of fat, cholesterol and bile acids, leading to reduced
hepatic and serum lipid levels.
Phytic acid has been reported to reduce the activity of
porcine pancreatic lipase in vitro (Knuckles et al. 1989)
and pancreatic lipase activity in the small intestine of
broilers (Liu et al. 2009), implying that the reduced fat
digestibility by phytic acid could partly be a result of
reduced activity of lipase. Phytic acid could also reduce
fat digestibility and absorption by binding bile acids via
divalent cations such as calcium to form insoluble phytic
acid-mineral-bile acids complexes, thereby reducing fat
digestion and absorption, and bile acids re-absorption.
However, this hypothesis needs to be tested.
With regards to protein digestibility, phytic acid has
been shown to reduce amino acid digestibility and to
increase the endogenous losses of nitrogen and amino
acids (see Tables 2 and 3). This implies that phytic acid
can reduce energy digestibility partly by reducing the
digestibility and increasing the endogenous losses of the
nitrogen and amino acids. For instance, Cowieson et al.
(2008) reported that an increase in the concentration of
phytic acid in a diet for broilers from 0.85 to 1.45%
resulted in an endogenous protein loss equivalent to
23 kcal kg1 of DM intake.
CONCLUSIONS
It can be concluded that phytic acid reduces animal
performance partly through reduced digestibility and
increased endogenous gut secretion of nutrients includ-
ing amino acids and minerals. The phytic acid-induced
reduction in nutrient digestibility is through binding to
nutrients, the digestive enzymes or both. The increased
endogenous losses of amino acids due to phytic acid
might be as a result of the increased secretion and
reduced re-absorption of digestive enzymes and mucins
in the gastrointestinal tract. Similarly, the increased
endogenous losses of divalent cations such as calcium
and zinc due to phytic acid might be as a result of the
increased secretion and reduced re-absorption of the
mineral-bicarbonates and digestive enzyme co-factors
in the gastrointestinal tract. The increased endogenous
loss of sodium due to phytic acid could mainly be due
to increased secretion as a result of increased secretion
of the mineral-bicarbonates; reduced re-absorption
due to reduced absorption of other nutrients, whose
absorption results in increased absorption of sodium;
or both. Overall, it appears that phytase, which is
added to pig and poultry feeds to improve phosphorus
availability, does not only improve phosphorus avail-
ability, but also alleviates the ant-nutritional effects of
phytic acid.
Further research is suggested to determine the effect
of dietary phytic acid on pig performance, true absorp-
tion of minerals in pigs and poultry, and true amino acid
and monosaccharide absorption and endogenous nutri-
ent losses in pigs. Also, further research is suggested to
determine the effect of dietary phytic acid on: (i)
activities of digestive enzymes (other than pepsin) and
gastrointestinal mucin secretions in pigs; (ii) secretion of
pancreatic mineral and digestive enzymes in the gastro-
intestinal tract of pigs and poultry to confirm whether
the increased ileal endogenous flow of sodium is due to
 WOYENGO AND NYACHOTI * ANTI-NUTRITIONAL EFFECTS OF DIETARY PHYTIC ACID 19
increased pancreatic mineral-bicarbonate secretion; (iii)
biliary secretions and absorption of bile acids in the
gastrointestinal tract of pigs and poultry; (iv) intestinal
histomorphology and synthesis of nutrient transporter
proteins in pigs and poultry; and (v) energy expenditure
in the gastrointestinal tract of pigs and poultry relative
to the whole body to establish whether the increased
endogenous secretion of nutrients increases intestinal
energy expenditure.
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 20 CANADIAN JOURNAL OF ANIMAL SCIENCE
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