Pakistan Journal of Nutrition 9 (8): 827-832, 2010
© Asian Network for Scientific Information, 2010
Corresponding Author: K.E. Akande, Animal Production Programme, School of Agriculture, Abubakar Tafawa Balewa University,
Bauchi, Bauchi State, Nigeria 827
Major Antinutrients Found in Plant Protein Sources: Their Effect on Nutrition
K.E. Akande , U.D. Doma , H.O. Agu and H.M. Adamu
1 1 2 3
Animal Production Programme, School of Agriculture,
Abubakar Tafawa Balewa University, Bauchi, Bauchi State, Nigeria
Department of Food Science and Technology, Federal Polytechnic, Bauchi, Bauchi State, Nigeria
Chemistry Programme, School of Science, Abubakar Tafawa Balewa University,
Bauchi, Bauchi State, Nigeria
Abstract: Compounds or substances which act to reduce nutrient intake, digestion, absorption and utilization
and may produce other adverse effects are referred to as antinutrients or antinutritional factors. Seeds of
legumes and other plant sources contain in their raw state wide varieties of antinutrients which are potentially
toxic. The major antinutrients includes: toxic amino acids, saponins, cyanogenic glycosides, tannins, phytic
acid, gossypol, oxalates, goitrogens, lectins (phytohaemagglutinins), protease inhibitors, chlorogenic acid
and amylase inhibitors. These antinutrients pose a major constraint in the use of plant protein sources in
livestock feeds without adequate and effective processing. The level or concentration of these anitnutrients
in plant protein sources vary with the species of plant, cultivar and post-harvest treatments (processing
methods). This paper reviews the nutritional effect of major antinutrients present in plant protein sources.
Key words: Antinutrients, plant protein, legumes
Antinutrients or antinutritional factors may be defined as
those substances generated in natural feedstuffs by the
normal metabolism of species and by different
mechanisms (for example inactivation of some
nutrients, diminution of the digestive process or
metabolic utilization of feed) which exerts effect contrary
to optimum nutrition. Being an antinutritional factor is not
an intrinsic characteristic of a compound but depends
upon the digestive process of the ingesting animal.
Trypsin inhibitors, which are antinutritional factors for
monogastric animals, do not exert adverse effects in
ruminants because they are degraded in the rumen
(Cheeke and Shull, 1985). Many plant components have
potential to precipitate adverse effects on the productivity
of farm livestock. These compounds are present in the
foliage and seeds of virtually every plant that is used in
practical feeding (D’Mello, 2000).
Nutritional effect of major antinutrients in plant protein
sources: The major antinutrients mostly found in plant
protein sources are toxic amino acids, saponins,
cyanogenic glycosides, tannins, phytic acid, gossypol,
oxalates, goitrogens, lectins (phytohaemagglutinins),
protease inhibitors, chlorogenic acid and amylase
Toxic amino acids: A wide range of toxic non-protein
amino acids occur in the foliage and seeds of plants.
These toxic non-protein amino acids appear to play a
major role in determining the nutritional value of a
number of tropical legumes (D’Mello, 1982). It has been
proposed that these amino acids act antagonistically
towards certain nutritionally important amino acids
(Liener, 1980). Fowden (1971) suggested that the
metabolic pathways culminating in the synthesis of
certain non-protein amino acids might reflect subtle
alteration in the genome responsible for directing the
formation of crucial amino acids. Bell (1971) reported
that while non-protein amino acids function primarily as
storage metabolites, they may also provide an adaptive
advantage to the plants, for example to render the plant
less susceptible to attack by various animals and lower
plants. Some of these toxic amino acids includes;
djenkolic acids, mimosine and canavanine.
Mimosine, a toxic non-protein amino acid structurally
similar to tyrosine, is contained in the legume Leucaena
leucocephala (D’Mello and Acamovic 1989; D’Mello,
2000). Mimosine has been proven effective in defleecing
sheep and goats (Jacquemet et al.,1990; Luo et al.,
2000). Mimosine a pyridoxal antagonist, which inhibits
DNA replication and protein synthesis; thus, it may elicit
defleecing by arresting cell division in the follicle bulb
(Reis, 1979). In monogastric animals, mimosine
causes poor growth, alopecia and reproductive
problems. Levels of Leucaena meal above 5-10% of the
diet for swine, poultry and rabbits generally result in poor
The major symptoms of toxicity in ruminants are poor
growth, loss of hair and wool, lameness, mouth and
Pak. J. Nutr. 9 (8): 827-832, 2010
oesophageal lesions, depressed serum thyroxine levelCyanogenic glycosides: Some legumes like linseed,
and goitre. Some of these symptoms may be due tolima bean, kidney bean and the red gram contain
mimosine and others to 3, 4-dihydroxypyridine, acyanogenic glycosides from which Hydrogen Cyanide
metabolite of mimosine in the rumen (Jones and(HCN) may be released by hydrolysis. Some cultivars of
Hegarty, 1984). Phaseolus lunatus (lima bean) contain a cyanogenic
Djenkol beans (Pithecolobium lubatum) whenglycoside called phaseolutanin from which HCN is
ingested sometimes lead to kidney failure which isliberated due to enzyme action, especially when tissues
accompanied by the appearance of blood and whiteare broken down by grinding or chewing or under damp
needle-like clusters in the urine. The clusters areconditions (Purseglove, 1991). Hydrolysis occurs rapidly
sulphur-containing amino acids known as djenkolicwhen the ground meal is cooked in water and most of
acids which are present in the bean in the free state, tothe liberated HCN is lost by volatilization. HCN is very
the extent of 1-4%. This toxic amino acid is structurallytoxic at low concentration to animals. HCN can cause
similar to cystine, but it is not degraded in the animaldysfunction of the central nervous system, respiratory
body. Due to its insolubility it crystallizes out in the kidney failure and cardiac arrest (D’Mello, 2000).
tubules and escapes through urine (Enwere, 1998).
The toxic, non-protein amino acid, canavanine, occursTannins: Tannins are water soluble phenolic
widely in unbound form in various legume plants of thecompounds with a molecular weight greater than 500
sub-family Papillonoideae (Bell et al., 1978) anddaltons. They have the ability to precipitate proteins from
abundantly in jack bean (Canavalia ensiformis (L). DC),aqueous solution. There are two different groups of
constituting up to 63 g/kg dry weight of the seed (Ho and tannins:- hydrolyzable tannins and condensed tannins.
Shen, 1966). Canavanine, a structural analogue ofCondensed tannins are widely distributed in
arginine, was first isolated from jackbean by Kitagawaleguminous forages and seeds. Cattle and sheep are
and Tomiyama (1929). sensitive to condensed tannins, while goats are more
Canavanine is believed to exert its toxic influence by resistant (Kumar, 1983; Kumar and Horigome, 1986;
virtue of its structural similarity with the nutritionallyKumar and Vaithiyanathan, 1990; D’Mello, 2000).
indispensable amino acid, arginine. Canavanine mayTannins may form a less digestive complex with dietary
antagonize arginine and interfere with Ribonucleic Acidproteins and may bind and inhibit the endogenous
(RNA) metabolism (Rosenthal, 1982). Canavanine hasprotein, such as digestive enzymes (Kumar and Singh,
been demonstrated to reduce feed intake of non-1984). Tannin-protein complexes involve both hydrogen-
ruminants but this was observed only at the equivalentbonding and hydrophobic interactions. The precipitation
of about 300 g/kg dietary level of raw jackbeanof the protein-tannin complex depends upon pH, ionic
(Tschiersch, 1962). strength and molecular size of tannins. Both the protein
Saponins: Saponins are a heterogeneous group ofthe precipitate increase with increase in molecular size
naturally occurring foam-producing triterpene orof tannins (Kumar and Horigome, 1986). However, when
steroidal glycosides that occur in a wide range of plants, the molecular weight exceeds 5,000 daltons, the tannins
including pulses and oil seeds such as kidney bean,become insoluble and lose their protein precipitating
chickpea, soybean, groundnut, lupin and sunflower capacity and degree of polymerization becomes
(Liener, 1980; Price et al., 1987; Jenkins and Atwal,imperative to assess the role of tannins in ruminant
1994). It has been reported that saponins can affect nutrition (Kumar, 1983; Lowry, 1990). Tannins have been
animal performance and metabolism in a number offound to interfere with digestion by displaying anti-trypsin
ways as follows: erythrocyte haemolysis, reduction ofand anti-amylase activity. Helsper et al. (1993) reported
blood and liver cholesterol, depression of growth rate,that condensed tannins were responsible for the testa-
bloat (ruminants), inhibition of smooth muscle activity,bound trypsin inhibitor activity of faba beans. Tannins
enzyme inhibition and reduction in nutrient absorptionalso have the ability to complex with vitamin B (Liener,
(Cheeke, 1971). Saponins have also been reported to1980). Other adverse nutritional effects of tannins have
alter cell wall permeability and therefore produce some been reported to include intestinal damage, interference
toxic effects when ingested (Belmar et al., 1999).with iron absorption and the possibility of tannins
Saponins have been shown to bind to the cells of theproducing a carcinogenic effect (Butler, 1989).
small intestine thereby affecting the absorption of
nutrients across the intestinal wall (Johnson et al., Phytic acid: Phytic acid occurs naturally throughout the
1986). plant kingdom and is present in considerable quantities
The effect of saponins on chicks have been reported towithin many of the major legumes and oilseeds. This
reduce growth, feed efficiency and interfere with theincludes soybean, rapeseed and cotton seed. Matyka et
absorption of dietary lipids, cholesterol, bile acids andal. (1993) reported that about 62-73% and 46-73% of the
vitamins A and E (Jenkins and Atwal, 1994). total phosphorus within cereal grains and legume
precipitation and incorporation of tannin phenolics into
Pak. J. Nutr. 9 (8): 827-832, 2010
seeds being in form of organically bound phytiningest considerable amounts of high-oxalate plants
phosphorus, respectively. As phytic acid accumulates inwithout adverse effects, due principally to microbial
storage sites in seeds, other minerals apparentlydecomposition in the rumen (Oke, 1969).
chelates to it forming the complex salt phytate (Erdman,The hulls of sesame seeds contain oxalates and it is
1979). Studies by Martinez (1977) revealed that inessential that meals should be completely decorticated
oilseeds, which contain little or no endosperm, thein order to avoid toxicities (McDonald et al., 1995).
phytates are distributed throughout the kernel foundChemical analysis carried by Alabi et al. (2005) on locust
within subcellular inclusions called aleurone grains orbean seeds revealed that the testa of locust bean seeds
protein bodies. had the highest concentration of oxalate (4.96 mg/100 g)
Whole soybeans have been reported to contain 1-2%followed by the pulp (3.40 mg/100 g) and the cotyledon
phytic acids (Weingartner, 1987; Osho, 1993). The major (1.15 mg/100 g). Olomu (1995) reported that pigeon pea
part of the phosphorus contained within phytic acid arecontains about 0.38% oxalic acid. Oxalic acid binds
largely unavailable to animals due to the absence of thecalcium and forms calcium oxalate which is insoluble.
enzyme phytase within the digestive tract of monogastric Calcium oxalate adversely affects the absorption and
animals. Nwokolo and Bragg (1977) reported that in theutilization of calcium in the animal body (Olomu, 1995).
chicken there is a significant inverse relationship
between phytic acid and the availability of calcium,Goitrogens: Goitrogenic substances, which cause
magnesium, phosphorus and zinc in feedstuffs such asenlargement of the thyroid gland, have been found in
rapeseed, palm kernel seed, cotton seed and soybeanlegumes such as soybean and groundnut. They have
meals. Phytic acid acts as a strong chelator, forming been reported to inhibit the synthesis and secretion of
protein and mineral-phytic acid complexes; the net result the thyroid hormones. Since thyroid hormones play an
being reduced protein and mineral bioavailabilityimportant part in the control of body metabolism their
(Erdman, 1979; Spinelli et al., 1983; Khare, 2000). Phytic deficiency results in reduced growth and reproductive
acid is reported to chelate metal ions such as calcium,performance (Olomu, 1995). Goitrogenic effect have
magnesium, zinc, copper, iron and molybdenum to formbeen effectively counteracted by iodine supplementation
insoluble complexes that are not readily absorbed fromrather heat treatment (Liener, 1975).
gastrointestinal tract. Phytic acid also inhibits the action
of gastrointestinal tyrosinase, trypsin, pepsin, lipase and Lectins (phytohaemagglutinins): Phytohaemagglutinins
"-amylase (Liener, 1980; Hendricks and Bailey, 1989;or lectins are glycoproteins widely distributed in
Khare, 2000). Erdman (1979) stated that the greatestlegumes and some certain oil seeds (including
effect of phytic acid on human nutrition is its reduction of soybean) which posses an affinity for specific sugar
zinc bioavailability. molecules and are characterized by their ability to
Gossypol: Gossypol is a naturally occurring polyphenolic (Pusztai, 1989). Lectins have the capability to directly
compound present in the pigment glands of cotton seedbind to the intestinal muscosa (Almeida et al., 1991;
(Gossypium spp). The average gossypol content varying Santiago et al., 1993), interacting with the enterocytes
from 0.4-2.4% within glanded cotton seeds to less thanand interfering with the absorption and transportation of
0.01% free gossypol within some low-gossypol cottonnutrients (particularly carbohydrates) during digestion
seed meals (Liener, 1980; Robinson and Brent, 1989;(Santiago et al., 1993) and causing epithelial lesions
Castaldo, 1995). Reduced lysine availability has beenwithin the intestine (Oliveira et al., 1989).
reported with cotton seed protein due to the ability ofAlthough lectins are usually reported as being heat-
gossypol to bind with the reactive epsilon amino grouplabile, their stability varies between plant species, many
of lysine during heat processing (Wilson et al., 1981;lectins being resistant to inactivation by dry heat and
Robinson, 1991; Church, 1991). The general symptomsrequiring the presence of moisture for more complete
of gossypol toxicity are depressed appetite, loss ofdestruction (Ayyagari et al., 1989; Poel et al., 1990;
weight, laboured breathing and cardiac irregularity.Almeida et al., 1991).
Death is usually associated with reduced oxygen-
carrying capacity of the blood, haemolytic effects on
erythrocytes and circulatory failure. Dietary gossypol also
causes olive-green discolouration of yolks in eggs
(Church, 1991; Olomu, 1995; McDonald et al., 1995).
Oxalates: Oxalates affects calcium and magnesium
metabolism and react with proteins to form complexes
which have an inhibitory effect in peptic digestion.
Ruminants, however unlike monogastric animals can
combine with carbohydrate membrane receptors
Protease inhibitors: Protease inhibitors are widely
distributed within the plant kingdom, including the seeds
of most cultivated legumes. Protease inhibitors have the
ability to inhibit the activity of proteolytic enzymes within
the gastrointestinal tract of animals (Liener and Kakade,
Trypsin inhibitor and chymotrypsin inhibitor are protease
inhibitors occurring in raw legume seeds. Protease
inhibitors are the most commonly encountered class of
Pak. J. Nutr. 9 (8): 827-832, 2010
antinutritional factors of plant origin. These inhibitorsprotein source, proper assessment of the type, nature
have been reported to be partly responsible for theand concentration of the antinutrients present in the
growth-retarding property of raw legumes. Theprotein source and also the bioavailability of nutrients to
retardation has been attributed to inhibition of proteinthe ingesting animal is necessary. Employing
digestion but there is evidence that pancreatic hyper-appropriate and effective processing techniques or
activity, resulting in increased production of trypsin andcombination of techniques could help reduce or
chymotrypsin with consequent loss of cystine andeliminate the adverse effects of these antinutritive
methionine is also involved (McDonald et al., 1995). constituents in plant protein sources and thereby
Trypsin inhibitors have been implicated in reducingimprove their nutritive value. Supplementation of some
protein digestibility and in pancreatic hypertrophyminerals, animo acids and vitamins could help reduce
(Liener, 1976). Trypsin inhibitors are polypeptides thator neutralize the negative effect of antinutritional factors
form well characterized stable complexes with trypsin on in plant protein sources for livestock nutrition. The
a one-to-one molar ratio, obstructing the enzymaticconcentration or level of the antinutritive constituents in
action (Carlini and Udedibie, 1997). Protease inhibitorsthese protein sources vary with the species of plant,
are inactivated by heat especially moist heat, because of cultivar and post-harvest treatments (processing
even distribution of heat (Bressani and Sosa, 1990;methods). Since antinutrients vary among plant cultivars,
Liener, 1995). therefore the use of genetically improved low-antinutritive
Chlorogenic acid: Sunflower meal contains high levelslivestock feeding.
of chlorogenic acid, a tannin like compound that inhibits
activity of digestive enzymes including trypsin,
chymotrypsin, amylase and lipase (Cheeke and Shull,
1985). Because chlorogenic acid is uncondensed and
non-hydrolyzable, its content of 1% or more of a total of
3-3.5% phenolic compounds in sunflower meal is not
reported in tannin assays. Chlorogenic acid is also a
precursor of ortho-quinones that occur through the
action of the plant enzyme polyphenol oxidase. These
compounds then react with the polymerize lysine during
processing or in the gut. Although the toxic effects of
chlorogenic acid can be counteracted by dietary
supplementation with methyl donors such as choline
and methionine. Chlorogenic acid is reported to be
readily removed from sunflower seeds using aqueous
extraction methods (Dominguez et al., 1993).
Amylase inhibitors: Amylase inhibitors are also known
as starch blockers because they contain substances
that prevent dietary starches from being absorbed by the
body. Starches are complex carbohydrates that cannot
be absorbed unless they are first broken down by the
digestive enzyme amylase and other secondary
enzymes (Marshall and Lauda, 1975; Choudhury et al.,
1996). Pigeon pea have been reported to contain
amylase inhibitors. These inhibitors have been found to
be active over a pH range of 4.5-9.5 and are heat labile.
Amylase inhibitors inhibit bovine pancreatic amylase but
fail to inhibit bacterial, fungal and endogenous amylase.
Pigeon pea amylase inhibitors are synthesized during
late seed development and also degraded during late
germination (Giri and Kachole, 1998).
Conclusion: The presence of antinutrients in plant
protein sources for livestock feeding is a major
constraint that reduces their full utilization. To be able to
justify the overall nutritional potential or value of any plant
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