Int. J. Food Ferment. Technol.

, 10(2): 101-111, December 2020

©2020 New Delhi Publishers. All rights reserved
DOI: 10.30954/2277-9396.02.2020.6

RESEARCH PAPER

Anti-Nutritional Factors in Food and Plant Crops

V.K. Joshi1*, Ghan Shyam Abrol2 and Amit Kumar Singh1
Department of Food Science and Technology, Dr Y S Parmar Universisty of Horticulture and Forestry Nauni, Solan, India
1

Department of Post-harvest Technology, Rani Lakshmi Bai Central Agricultural University, Jhansi, India
2

*Corresponding author: [email protected]

Paper No.: 248 Received: 15-09-2020 Revised: 22-11-2020 Accepted: 14-12-2020

ABSTRACT
Some of plants mainly contains toxic compound such as saponins, tannins, phytic acid, gossypol, lectins, protease
inhibitors, amylase inhibitor, goitrogens etc. which limits their consumption and also reduces nutrient bioavailability when
consumed. These factors are responsible for micronutrient malnutrition and mineral deficiencies. On the other hand they
act as self defense system to the food. These are categorized into different categories and causes various symptoms upon
consumption such as acute toxicity include respiratory distress, impaired body weight gain, anorexia, weakness, apathy
and in some cases even death after several days. But there are various traditional methods and improved technologies are
available, which can be used to reduce the levels of these anti-nutrient factors. Various processing methodologies such as
fermentation, germination, debranning, autoclaving, soaking, drying, irradiation etc. are used to reduce the anti-nutrient
contents in foods. The review is focused on some common recently important anti-nutritional factors (ANF) which were
of major concern.
Keywords: Anti-nutrients, Saponins, Trypsin Inhibitors, Cooking, Fermentation, Irradiation

The anti-nutritional factors (ANF) are the toxic feeds (Fig. 1). They are enzyme inhibitors (trypsin
compounds synthesized naturally in plant, microbes and chymotrypsin inhibitors, plasmin inhibitors,
and animals cause serious problems in human and elastase inhibitors), haemaglutinnins (concanavalin
animal nutrition. These are compounds which A, ricin), plant enzymes (urease, lipoxygenase),
reduces the nutrient utilization or food intake when cyanogenic glycosides (phaseolunatin, dhurrin,
consumed as foods (Soetan and Oyewole, 2009). The linamarin, lutaustralin), goitrogens (pro-goitrins,
presence of these endogenous compounds results in glucosinolates), oestrogens (flavones, genistein),
limiting use of food or feed stuff. saponins (soya sapogenin), gossypol from
Gossypium species e.g., cotton, tannins (condensed
The ANFs can be divided into two major categories.
and hydrolysable tannins), amino acid analogues
One category is of proteins (such as lectins and protease
(BOAA, DAP, mimosine, N-methyl-1- alanine),
inhibitors) which are sensitive to normal processing
alkaloids (solanine, chaconine), anti-metals (phytates,
temperatures and second is of other substances
oxalates), anti-vitamins (anti-vitamins A, D, E and
which are stable or resistant to these temperatures
B12) and favism factors.
and include among many others polyphenolic
compounds (mainly condensed tannins), non-protein How to cite this article: Joshi, V.K., Abrol, G.S. and Singh, A.K. (2020).
amino acids and galactomannan gums. According to Anti-Nutritional Factors in Food and Plant Crops. Int. J. Food Ferment.
Technol., 10(2): 101-111.
Aletor (1993) there are several ANFs that are very
Source of Support: None; Conflict of Interest: None
significant in plants used for human foods and animal
 Joshi et al.

foaming characteristic (Price et al. 1987). The word

is derived from Latin word sapo from soapwort
plant (Saponaria). They are either a choline steroid
or triterpenoid attached via C3 and an ether bond
to a sugar side chain. Most of the saponins occur as
insoluble complexes with 3-b-hydroxysteroids which
interact with bile acid and cholesterol, similarly,
saponin with either of iron, zinc, and calcium form
insoluble complexes (Oakenfull et al. 1989; Milgate
and Roberts, 1995; Rafińska et al. 2017). Soybean,
chickpea, faba bean, pea, lentil and peanuts are the
different sources of saponins. Erythrocytes lyse in
Fig. 1: Classification of anti-nutrients present in food saponin solution, therefore these compounds are
(Samtiya et al. 2020)
toxic when injected intravenously (Khalil and El-
Although these factors vary in their individual Adawy, 1994). Saponin content may be variable, even
effect, a large proportion of them can be destroyed among the same species of edible beans, because of
or inactivated simply by heat treatment processes variations in cultivars, varieties, locations, irrigation
during cooking. Unfortunately, detailed toxicological condition, type of soil, climatic conditions, and
studies have not been performed on the majority of year during which they are grown (Price et al. 1987;
these anti-nutritional factors. Broadly speaking, their Khokhar & Chauhan, 1986; Fenwick & Oakenfull,
presence in untreated foodstuffs normally results in 1983; Gholami et al. 2014; Bianca et al. 2020). High
anorexia, reduced growth and poor food conversion concentrations impart a bitter taste and astringency
efficiency when used at high dietary concentrations. in dietary plants causes its limited use as food
Most of legumes are under-used because of the (Liener, 1994). The anti-nutritional effects of saponins
presence of anti-nutritional compounds, such as have been mainly studied using alfalfa saponins.
enzyme (trypsin, chymotrypsin, alpha-amylase) Saponins cause hypocholesterolaemia by binding
inhibitors, phytic acid, flatulence factors, saponins cholesterol, making it unavailable for absorption.
and toxic factors (Lyimo et al. 1992). Root crops, in They also cause haemolysis of red blood cells and
common with most plants, contain small amounts are toxic to rats (Johnson et al. 1986). Saponins have
of potential toxins and antinutritional factors such wide applications in beverages, confectionery and
as trypsin inhibitors. Apart from cassava that pharmaceutical products (Sparg et al. 2004; Petit et al.
has cyanogenic glucosides, cultivated varieties of 1995; Uematsu et al. 2000).
most edible tubers and roots do not contain any
serious toxins. Thus, wild species must be properly
processed before their consumption. These wild
species, however, are useful reserves in times of
famine or food scarcity. Local people are aware of
their potential risks in their use and have developed
indigenous techniques to detoxify such crops before
consumption.

SAPONINS
Saponins are glycosides present in plants and due
to hydrophobic nature of aglycone and hydrophilic Fig. 2: Presence of total saponins content in commonly
nature of sugar chains results in distinctive used spices (Sirohi et al. 2009)

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 Anti-Nutritional Factors in Food and Plant Crops

Soaking, blanching, cooking and other treatments growth retardation. Trypsin (protese inhibitor) causes
reduce the saponins content. Cooking of legumes pancreatic enlargement and growth depression
reduces the amount of saponins by 7–53% (Jood (Aletor and Fetuga, 1987). Cooking and germination
et al. 1986; Sharma and Sehgal, 1992). Cooking of seem to be good procedures to improve the quality
chickpea and black gram and of faba beans lowered of lentil flour from the nutritional point of view,
the saponin content by 7–17% and 35%, respectively despite the fact that a large variation on the effects of
(Kataria et al. 1988). processing, related to the different legume varieties,
has been observed (Vidal et al. 1994).
TRYPSIN INHIBITOR
Legumes are low cost source of protein in the diet Table 1: Different treatments to reduce the Trypsin inhibitor
(Borade et al., 1984). But another side they also
contains some anti-nutritional factors like trypsin Treatments Effect References
which is injurious to human health (Soetan and Thermal Pressure cooking reduces Habiba, 2002
Oyewole, 2009). Trypsin inhibitior strappingly treatments trypsin inhibitors by 50%
prevent the activity of key pancreatic enzymes while microwaving 25%

trypsin and chymotrypsin thus reducing digestion Chickpeas needed Alajaji and El‐
and absorption of proteins by the formation of minimum 90 minutes of Adawy, 2006
boiling or 35 minutes of
different complexes (Gemede and Ratta, 2014). These
autoclaving or 15 minutes
are classified in 2 families as per their molecular size: of microwaving
Kunitz (KTIs), having molecular weights around 20
Lentils, require only 1 Embaby, 2010a;
kDa and Bowman‐Birk (BBTIs) around 8 kDa (Avilés‐
min roasting at 80 °C to Ma and other
Gaxiola et al. 2018). The tripsin inhibitor content of reduce up to 95.6% trypsin 2011
legumes increased during germination. The intake of inhibitors and also total
trypsin inhibitor with the diet causes a rise of fecal tannins and phytic acid.
nitrogen loss (Combs et al. 1967). 4 h soaking followed by Pedrosa and
cooking at 95 °C for 1 h the others 2015; Shi
trypsin inhibition of pea, and others 2017
lentil, faba bean, chickpea,
and common bean
reduced 81.25%, 100%,
100%, 88.37%, and 93.7 %,
respectively
Extrusion Extrusion cooking after Adamidou and
pre-treatment at different others 2011
high temperatures to
the legumes field pea,
chickpea and faba bean
*Trypsin inhibition activity of Soyabean is 94.1 U/mg (Yalcin and flour resulted in reductions
Basman, 2015) of up to 53.7%, 91.8%, and
58.9%, respectively
Fig. 3: Trypsin inhibitor activity in different legumes
(Avilés‐Gaxiola et al. 2018) Soybeans and lentil, Žilić and others
extrusion at 150 °C found 2012; Rathod and
Trypsin inhibitor may have different modes of action, to be enough to inactivate Annapure 2016
such as the antifibrinolytic or antithrombinogenic trypsin inhibitors activity
or antiproteolytic or a promoter of conversion of up to 95%
methionine to cysteine manifesting ultimately in

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 Joshi et al.

Ultrasound Ultrasound waves 30 kHz Huang and CYANOGENS (CNglcs)

for 20 min in soybeans others 2008
Cyanogenic glucosides are phytoanticipins of more
decreases 55% of Kunitz
Trypsin inhibitor than 60 different types and known to be present in
High Protein digestibility was Linsberger‐ more than 2500 plant species (Conn, 1980; Bak et al.
hydrostatic improved by 4.3% and Martin and 2006). Generally, CNglcs acts as defense mechanism
pressure 8.7% in split peas and others 2013 to plants which resulted in release of toxic hydrogen
white beans
cyanide (HCN), an aldehyde or ketone and glucose
Infrared IR exposure completely Bellido et al. 2006
compounds (Nahrstedt, 1996; Vetter, 2000). The lethal
radiation (IR) inactivates trypsin
inhibitor and lipoxygenase dose of cyanide for vertebrates lies in the range of
enzymes in soybean 35–150 µmol/kg (Zagrobelny et al. 2008). Generally,
Soaking Simplest way of El‐Adawy et al. cyanogenic glycoside is considered as not toxic on its
inactivation of trypsin 2000; Mumba et own (Bolarinwa et al. 2016).
inhibitor activities al. 2004
Germination Germination reduces Sangronis and A cyanogenic food of particular economic importance
trypsin inhibitor Machado 2007 is cassava, which is also known by the names manioc,
Fermentation Solid‐State Fermentation in Gao et al. 2013 yuca and tapioca. Cassava roots have been reported
A. oryzae over a period of 5
to contain cyanide content of 10–500 mg/kg of dry
days reduces 89.2% trypsin
activity matter (Siritunga and Sayre, 2003). Amygdalin is
the cyanogenic glycoside responsible for the toxicity
MIMOSINE found in the range of 8.84–48.33 mg/g in Cassava
Mimosine [ß-N-(3-hydroxy-4-pyridone)-a-amino- (Bolarinwa et al. 2016) and also in many species
propionic acid], a non-protein amino acid structurally of Rosaceae, such as bitter almonds, peaches and
resembles to tyrosine and found in the genera Leucaena apricots (Bolarinwa et al. 2014). Sweet almonds are low
and Mimosa. The mimosine contents in different in amygdalin as a result of breeding processes. The
parts varies differently as in seeds 4 to 5%, in roots lethal dose of HCN for cattle and sheep is 2.0-4.0mg
1 to 1.5% and in shoot 1 to 12% similarly old stems per kg body weight. The lethal dose for cyanogens
have the smallest and growing tips contains largest would be 10-20 times greater because the HCN
amounts (Jones and Lowry, 1984). The mechanism of comprises 5-10% of their molecular weight (Conn,
action of mimosine in producing its effect, however, 1979). Cyanide content of bamboo shoot ranged from
is not clear but it may act as an amino acid antagonist 1000 to 8000 mg/kg hydrogen cyanide (Ferreira et al.
or may complex with pyridoxal phosphate, leading 1995). For poisoning, forage containing this amount
to disruption of catalytical action of B6-containing of cyanogens would have to be consumed within a
enzymes such as trans-aminases, or may complex few minutes and simultaneous HCN production
with metals such as zinc (Hegarty, 1978). The main
would have to be rapid. Recorded accounts of
symptoms of toxicity in ruminants are poor growth,
livestock poisoning by cyanogenic plants show that
loss of hair and wool, swollen and raw coronets
such situations do arise. Cyanogens have also been
above the hooves, lameness, goitre etc. (Jones and
Hegarty, 1984). Moist heat at 70 to 100°C reduce 50% suspected to have teratogenic effects (Keeler, 1984).
mimosine content (Akbar and Gupta, 1985) and dry Post-harvest wilting of cyanogenic leaves may reduce
heat treatment at 100°C reduced 17 to 19% mimosine the risk of cyanide toxicity. Animals suffering from
content whereas autoclaving reduced 19 to 23% (Mali cyanide must be immediately treated by injecting
et al. 1990). a suitable dose of sodium nitrate and sodium
thiosulphate.

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 Anti-Nutritional Factors in Food and Plant Crops

Table 2: Different treatments to reduce the cyanogens The content of total oxalate, soluble oxalate, calcium
oxalate and free calcium for various aroids as
Treatments Effect References
summarized in Table 3. Holloway et al. (1989) found
Soaking Soaking of cassava root Kemdirimi et al.
that the stems of giant swamp taro, elephant foot
for72 h reduces 90% of 1995
cyanide content
yam, skin of giant taro and taro leaves contained
Soaking of apricot Tuncel et al. 1995
about 400 mg/l00 g fresh weight of calcium oxalate,
kernels at 20°C about 10 times the amount present in taro Xanthosoma
significantly reduces and Colocasia.
amygdalin
Fermentation Fermentation of cassava Kemdirimi et al., Table 3: Content of calcium oxalates and calcium (mg/100 g
pulp or dough for 4–5 1995 fresh weight) in roots and stems of tropical root crops (standard
days has been reported deviation in parentheses)
to decrease its total
cyanide by 52–63% Total Soluble Calcium Total
Fermentation of Igbadul et al. 2014 Root crop and source
oxalate Oxalate Oxalate Ca
cocoyam flour reduces 1. taro, Colocasia esculenta
cyanide by 98.6%
(a) Fiji croms, mean of 5
Fermentation results in Prasad and Dhanya, 65 (19) 35 (4) 43 23 (5)
Cvs.
reduction of 84.6% in 2011
(b) Fiji, suckers, mean of 2
the cyanide content in 60 (1) — — 14 (2)
Cvs.
sorghum leaves
2. taro, X. sagittifolium
Storage Cyanide content Onabolu et al. 2002
tonga corms, mean of 3 60 (30) 44 (22) 23 6 (1)
decrease by 50–64%
Cvs.
in the cassava product
when stored for 4 weeks 3. giant taro, Alocasia
at room temperature (a) Western Samoa, mean
38 (18) 17 (9) 31 26 (5)
of 6 Cvs.
OXALATES (b) Western Samoa mean of
30(8) — — 28 (9)
6 Cvs.
Oxalates were found in various families including 4. giant swamp taro,
aroid / arum family e.g., Colocasia (taro) and Crytosperma
Xanthosoma (Calodium) (Noonan and savage, (a) Fed. States of
319 135
1999). Oxalic acid result of oxidative breakdown of Microneisa (Ponape), Mean 45 (39) 399
(77) (33)
carbohydrate and protein, accumulated in plants of 2 Cvs.
especially during dry conditions (Bressani et al. 1993). 300 219
(b) Kiribati, mean of 5 Cvs. — —
Reports of some tropical leafy vegetables revealed (218) (156)
that dry vegetables had higher oxalate concentrations Source: Holloway et al. (1989).
than the fresh vegetables (Aletor and Adeogun, 1995).
Raw, unpeeled Colocasia (Colocasia esculenta) corm
Oxalate content in taro (Colocasia esculenta) contain
meal contained total oxalate 1234 mg/kg of dry
278-574 mg /100 g fresh weight, sweet potato (Ipomoea
matter. About a third of the oxalates were removed
batatas) 470 mg/100 g fresh weight and yam (Dioscorea
during peeling the outer skin of the corms, and when
alata) tubers 486- 781 mg/100 g dry were reported
peeling was combined with boiling, oxalate contents
(Holloway et al. 1989; Mosha et al. 1995). Oxalate
were reduced to 272 mg/kg. The data suggested
was widely distributed in plants in a readily water
that oxalate was a major factor contributing to
soluble form as potassium, sodium and ammonium
the antipalatability and antinutritive effect of raw
oxalate and as insoluble calcium oxalates (Fassett,
Colocasia corm. Shajeela et al. (2011) observed that
1973; Connor, 1977; Smith, 1982).
D. bulbifera var. vera content highest amount of total

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 Joshi et al.

free phenolics (2.20g/100g on dry wt. basis) and total Spinach, Rhubarb stalk, Beetroot, Broccoli, Carrot,
oxalate (0.78g/100g on dry wt. basis) in all species Parsnip) Savage et al. (2000). Freezing followed
of yam. Bhandari and Kawabata (2004) studied on by grilling reduces the availability of calcium as
oxalate content in four species of yam and found 76.7% of calcium was bound to oxalate to insoluble
that the D. deltoidea content highest amount of total oxalate form (Brogren and Savage, 2003). Similarly,
oxalate, soluble oxalate and calcium oxalate followed the oxalate content of spinach pasta was found to
by D. triphylla, D. versicolor and D. Bulbifera. contained highest amount of total (91.8 mg/100g
wet weight) and soluble oxalate (53.8 mg/100g wet
Singh et al., (1999) studied on biochemical composition
weight (Liebman and Okombo, 2009).
and nutririve value of promising collections of
elephant foot yam of five genotype like Am-1, Am-2,
GLYCOALKALOIDS
Am-3, Am-4, Am-5, and found that Am-5 contained
highest amount of oxalate (0.96 %) and Am-1 (0.58 %) Potatoes are good source of glycol-alkaloids alpha-
was the lowest. Chattopadhyay et al. (2010) studied solanine and alpha-chaconine, concentrated mainly
on quantitative and qualitative aspects of elephant in the flowers and sprouts (200 to 500 mg/100
foot yam corm and found that the genotype NDA- g). For food safety purposes, an upper limit for
4 contained highest amount of soluble oxalate (18.50 glycoalkaloid content of 20 mg per 100 g of potato is
mg/100g) while, Midnapur Finger was lowest in generally accepted. Concentrations of glycoalkaloids
amount (6.65 mg/100g). Oscarsson and Savage (2007) are 3 to 10 times greater in the peel than in the flesh.
studied on composition and availability of soluble and In bitter varieties, the alkaloid concentration can go
insoluble oxalates in raw and cooked taro (Colocasia upto 80 mg/100 g in the tuber as a whole and up to
esculenta var. Schott) leaves and found that young 150-220 mg/100 g in the peel. At these concentrations
taro leaves contained 589 mg total oxalate/100g fresh of solanine and other potato glycoalkaloids are toxic.
wt. while older taro leaves contained 443 mg total But because of higher temperature of decomposing
oxalate/100g fresh wt. and soluble oxates were 74% (243 °C) they remain unaffected at normal cooking.
of total oxalate content of the young and old leaves. To avoid toxic levels of glycoalkaloids, potato cultivar
They also studied on soluble and insoluble oxalate in selection is very important. However, improper
young and old leaves with treatment of fresh, baked postharvest handling conditions are the main cause
and baked with milk and found that the old leaves of toxic levels in potatoes. To keep glycoalkaloid
contained highest per cent of soluble and lowest per content low, store potatoes at lower temperatures,
cent of insoluble oxalate, while, old leaves baked with such as 7°C, keep potatoes away from light, market
milk contained lowest soluble and highest insoluble in opaque plastics films and paper bags, and rotate
per cent of oxalate. Frank et al., (2001) studied on frequently on retail displays.
oxalate, insoluble calcium and soluble oxalate in
Phaseolus vulgaris leaves and found that the adult leaf LECTINS
contained more amount of oxalte than the growing Lectins are proteins or glycoproteins with molecular
leaf on dry weight basis. weight of 10, 000–124, 000. Lectins, also referred to as
Hang et al. (2012) studied on effect of simple washing phyto-haemagglutinins, are glycoprotein compounds
processing methods on oxalate content of taro which have been shown to agglutinate red blood cells
petioles and found that petioles and leaves of cv. in vitro (Gatel, 1994). It is also defined as ‘proteins
Chia Voi contained lowest soluble and total oxalate or glycoproteins of non-immune origin with one or
in washed sample, and highest in raw sample. more binding sites per subunit, which can reversibly
Cooking at 45 minute at 15 psi pressure reduced the bind to specific sugar segments through hydrogen
total, soluble and insoluble content of all vegetables bonds and Van Der Waals interactions’ (Lis and
(silverbeet leaf, silverbeet stem, New Zeland spinach, Sharon, 1998). Lectin activity has been determined in

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 Anti-Nutritional Factors in Food and Plant Crops

more than 800 varieties of the legume family. Of the aqueous heat treatment of fully imbibed soy seeds at
total protein 2- 10 per cent legume seeds are lectins. 100°C for 10 min.
One of their most important characteristics is that
they prevent absorption of digestive end products GOSSYPOL
in the small intestine. In general, plant lectins are Gossypol is a phenolic compound first isolated in
most abundant in the seeds but they are also found 1899 and its name is derived from the genus scientific
in different vegetative tissues such as in roots, leaves, name (Gossypium) combined with the “ol” from
barks, flowers, bulbs and rhizomes (Ratanapo et al. phenol (Soto-Blanco, 2008). Gossypol is produced by
1998; Van Damme et al. 2000). Soybean, common pigment glands in cotton stems, leaves, seeds, and
bean, jack bean, chickpea, faba bean, pea, lentil and flower buds but the highest concentration is seen
peanut are good sources of lectins. Peas generally in seeds (Rogers et al. 2002; Kenar, 2006; Alexander
have higher lectin activities than faba beans, but both et al. 2008). General signs of acute toxicity include
show considerably lower amounts in comparison respiratory distress, impaired body weight gain,
to raw defatted soybeans. Effects of feeding lectin anorexia, weakness, apathy and in some cases even
containing diets include changes in gut immune death after several days (Fombad and Bryant, 2004).
function, reduced production of endocrine cells and The amount of free gossypol in the cottonseed can
gut hormones, interference with the bacterial ecology be quite variable. Many factors influence gossypol
in the gut lumen, and damage to mucosal cells (King et content such as: specie of cotton plant, climatic
al., 1983). Lectins have the capability to directly bind to conditions, soil conditions, fertilizer, etc. Since
the intestinal mucosa interacting with the enterocytes gossypol affects the heart, gossypol toxicity has
and interfering with absorption and transportation been manifested as two types of clinical syndromes
of nutrients (particularly carbohydrates) during particularly in young animals. Dietary gossypol can
digestion and causing epithelial lesions within the also bring about increased requirement, not only
intestine (Santiago et al. 1993, Oliveira et al. 1989). for lysine, but also for iron which it can chelate. At
The main toxic components in P. vulgaris are lectins, the physiological level, gossypol reduces oxygen
sugar-binding proteins which bind and agglutinate availability in the blood. One syndrome of sudden
red blood cells. The toxicity of lectins is characterized death (resembling a heart attack) has frequently been
by growth inhibition in experimental animals and reported in calves and lambs. Dietary free gossypol
diarrhoea, nausea, bloating and vomiting in humans. of up to 0.02-0.03% has been reported to cause death
But thorough cooking can destroy this to a safe limit. in growing pigs while poultry can tolerate fairly
Armour et al. (1998) reported a complete inactivation high dietary levels (Aletor and Onibi, 1990). These
of soy lectin and protease inhibitory activity by animals seem healthy, have good appetites and are
Table 4: Treatments to Reduce Gossypol Toxicity

Treatments Effect References

Heat treatment Roasting & Extrusion both reduces the Arieli,1998; Noftsger et al. 2000
gossypol content in milk
Irradiation Gamma irradiation & Electron beam Quintana et al. 2000; Sijun et al. 2012; Shawrang et al. 2011;
irradiation reduces gossypol content in Ebrahimi-Mahmoudabad and Taghinejad-Roudbaneh,
cottonseed meal 2011
Fermentation Fungus Aspergillus niger, Aspergillus oryzae, Barros et al. 2002; Yildirim et al. 2003; Lim and Lee, 2011
Candida tropicalis, Saccharomyces cerevisiae,
Geotrichum candidum found to be very effective
in reducing Gossypol Toxicity

107
 Joshi et al.

often one of the best ones in the group, but are found Armour, J.C., Perera, R.L.C., Buchan, W.C. and Grant, G. 1998.
dead. If cotton seed or cotton seed meal is bought in Protease inhibitors and lectins in soya beans and effects of
aqueous heat-treatment. J. Sci. Food Agric., 78: 225–231.
bulk, it would be worthwhile to have it tested before
Avilés‐Gaxiola, S., Chuck‐Hernández, C. and Serna Saldívar,
feeding it. Depending on the level of free gossypol,
S.O. (2018), Inactivation Methods of Trypsin Inhibitor in
the cottonseed could be utilized in the best interest of Legumes: A Review. Journal of Food Science, 83: 17-29.
the livestock. Bak, S., Paquette, S.M., Morant, M., Rasmussen, A.V., Saito,
S., Bjarnholt, N., Zagrobelny, M., Jørgensen, K., Hamann,
SUMMARY T., Osmani, S., Simonsen, H.T., Pérez, R.S., van Hesswijck,
T.B., Jørgensen, B. and Møller, B.L., 2006. Cyanogenic
Antinutritional factors like saponin, trypsin glycosides; a case study for evolution and application of
inhibitors etc. are the defense mechanism of plants, cytochromes P450. Phytochem. Rev., 5: 309–329
yet these cause many ill effects when consumed. Barros, M.M., Lim, C. and Klesius, P.H. 2002. Effect of
Antinutritional factor is a serious problem in wild soybean meal replacement by cottonseed meal and
species while most of cultivated species are devoid iron supplementation on growth, immune response
and resistance of channel catfish (Ictalurus puctatus) to
of it. Beside these, many processing practices are
Edwardsiella ictaluri challenge. Aquaculture, 207(3-4): 263–
developed in these days to avoid the ill effect of 279.
antinutritional factors. Since, the antinutrional factors Bhandari, M.R. and Kawabata, J. 2004. Assessment of
are defense mechanism of plants so these need to be antinutritional factor and bioavailability of calcium and
exploited for the benefits of human use. zinc in wild yam (D. spp) tubers of Nepal. Food Chemistry,
85: 281-287.
REFERENCES Bianca Ribeiro, Elia Lacchini, Keylla U. Bicalho, Jan Mertens,
Philipp Arendt, Robin Vanden Bossche, Gabriela Calegario,
Akbar, M.A. and Gupta P.C. 1985. Subabul (Leucaena
Lore Gryffroy, Evi Ceulemans, Julia Buitink, Alain
leucocephala) as a source of protein supplement for buffalo
Goossens and Jacob Pollier. 2020. A Seed-Specific Regulator
calves. Indian Journal of Animal Sciences, 55: 54-58.
of Triterpene Saponin Biosynthesis in Medicago truncatula.
Aletor, V.A. 1993. Allelochemicals in plant foods and The Plant Cell, 32(6): 2020-2042.
feeding stuffs. Part I. Nutritional, biochemical and
Bolarinwa, I.F., Oke, M.O., Olaniyan, S.A. and Ajala, A.S. 2016.
physiopathological aspects in animal production. Veterinary
A Review of Cyanogenic Glycosides in Edible Plants. DOI:
and Human Toxicology, 35(1): 57-67.
10.5772/64886
Aletor, V.A. and Fetuga, B.L. 1987. Pancreatic and intestinal
Bolarinwa, I.F., Olaniyan, S.A., Olatunde, S.J., Ayandokun, F.T.
amylase (EC 3.2.1.1) in the rat fed haemagglutinin extract.
and Olaifa, I.A. 2016. Effect of processing on amygdalin
II Evidence of impaired dietary starch utilization. J. Anim.
and cyanide contents of some Nigerian Foods. J. Chem.
Physiol. Anim. Nutr., 57(3): 113-117.
Pharm. Res., 8(2): 106‐113.
Aletor, V.A. and Onibi, O.E. 1990. The use of oyster shell as
Bolarinwa, I.F., Orfila, C. and Morgan, M.R.A. 2014. Amygdalin
calcium supplement. Part I. Effect on the utilization of
content of seeds kernels and food products commercially‐
gossypol-containing cottonseed cake by the chicken. Die.
available in the UK. Food Chem., 152: 1333‐139.
Nahrung. Food, 34(4): 311-319.
Borade, V.P., Kadam, S.S. and Salunke, D.K. 1984. Changes in
Aletor, V.A. and Adeogun, O.A. 1995. Nutrient and
phytate phosphorus and minerals during germination and
antinutrient component of some tropical leafy vegetables.
cooking of horse gram and moth bean. Qual. Plant Foods
Food Chem., 53: 375-379.
Hum. Nutr., 34: 151.
Alexander, J., Benford, D., Cockburn, A. et al. 2008. Gossypol
Bressani, R. 1993. Amaranth. In: Encyclopaedia of food science,
as undesirable substance in animal feed. EFSA Journal, 908:
food technology and nutrition. Eds. Macrae R, Rabinson
1–55.
RK, Sadler, MJ. London. Academic Press, pp. 135-140.
Arab, A.A.A. and Salem, F.M.A. 2010. Nutritional quality
Brogren, M. and Savage, G.P. 2003. Bioavailability of soluble
of Jatropha curcas seeds and effect of some physical and
oxalate from spinach eaten with and without milk products.
chemical treatments on their anti-nutritional factors.
Asia Pacific J. Clin. Nutr., 12(2): 219-224.
African Journal of Food Science, 4(3): 93-103.
Chattopadhyay, A., Saha, B., Pal, S., Bhattacharya, A. and Sen,
Arieli, A. 1998. Whole cottonseed in dairy cattle feeding: a
H. 2010. Quantitative and qualitative aspects of elephant
review. Animal Feed Science and Technology, 72(1-2): 97–110.
foot yam. International Journal of Vegetable Science, 16: 73-84.

108
 Anti-Nutritional Factors in Food and Plant Crops

Combs, G. E., Conness, R.G., Berry, T.H. and Wallace, Holloway, W.D., Argall, M.E., Jealous, W.T., Lee, J.A. and
H.D. 1967. Effect of raw and heated soybeans on gain, Bradbury, J.H. 1989. Organic acids and calcium oxalate in
nutrient digestibility, plasma amino acids and other blood tropical root crops. J. Agic. Food Chem., 37: 337-341.
constituents of growing swine. J. Anim. Sci. 26: 1067-1071. Igbadul, B.D., Amoye, J. and Twadue, I. 2014. Effect of
Conn, E.E. 1979. Cyanide and cyanogenic glycosides. In: Hcr- fermentation on the proximate composition, antinutritional
bi vores: Their interaction with secondary plant metabolites. factors and functional properties of cocoyam (Colocasia
Rosenthal, G.A. Janzen, D.H. (eds.) A.P., New York pp. 387- esculenta) flour. Afr. J. Food Sci. Technol., 5(3): 67‐74.
412. Johnson, I.T., Gee, J.M., Price, K., Curl, C. and Fenwick, G.R.
Conn, E.E. 1980. Cyanogenic compounds. Ann. Rev. Plant 1986. Influence of saponin on gut permeability and active
Physiol., 31: 433–451. nutrient transport in vitro. J. Nutr., 116: 2270-2277.
Connor, H.E. 1977. The poisonous plants in New Zealand: Jones, R.J. and Hegarty, M.P. 1984. The effect of different
Wallington, New Zealand Government Printer, pp. 134. proportion of Leucaena leucocephala in the diet of cattle
on growth, feed in­ take, thyroid function and urinary
Ebrahimi-Mahmoudabad S.R. and Taghinejad-Roudbaneh
excretion of 3-hydroxy-4 (lH)-pyridone. Australian Journal
M. 2011. Investigation of electron beam irradiation effects
of Agriculture Research, 35: 317.
on anti-nutritional factors, chemical composition and
digestion kinetics of whole cottonseed, soybean and canola Jood, S., Chauan, B.M. and Kapoor, A.C. 1986. Saponin content
of chickpea and black gram: varietal differences and effects
seeds. Radiation Physics and Chemistry, 80(12): 1441–1447.
of processing and cooking methods. J. Food Agric., 37: 1121–
Fassett, D.W. 1973. Oxalates: In toxicants occurring naturally 1124.
in Foods. Washington D.C. National Acad. Scie., pp. 346-362.
Kataria, A., Chauhan, B.M. and Gandhi, S. 1988. Effect of
Fenwick, D.E. and Oakenfull, D.G. 1983. Saponin content of domestic processing and cooking on the antinutrients of
food plants and some prepared foods. J. Sci. Food Agric., black gram. Food Chem., 30: 149–156.
34: 186–191.
Keeler, R.W. 1984. Teratogens in plants. Journal of Animal
Ferreira, V.L.P., Yotsuyanagi, K. and Carvalho, C.R.L. 1995. Science, 58: 1029-1039.
Elimination of cyanogenic compounds from bamboo shoots
Kemdirimi, O.C.C., Chukwu, O.A. and Anchinewhu, S.C.
Dendrocalamus giganteus Munro. Trop. Sci., 35: 342‐346. 1995. Effect of traditional processing of cassava on the
Fombad, R.B. and Bryant, M.J. 2004. An evaluation of the use cyanide content of gari and cassava flour. Plant Foods Hum.
of cottonseed cake in the diet of growing pigs. Tropical Nutr., 48: 335‐339.
Animal Health and Production, 36(3): 295–305. Kenar, J.A. 2006. Reaction chemistry of gossypol and its
Frank, E.Z., Honow, R. and Hesse, A. 2001. Calcium and derivatives. Journal of the American Oil Chemists’ Society,
oxalate content of the leaves of Phaseolus vulgaris at 83(4): 269–302.
different calcium supply in relation to calcium oxalate Khalil, A.H. and El-Adawy, T.A. 1994. Isolation, identification
crystal formation. J. Plant Physiol., 158: 139–144. and toxicity of saponin from different legumes. Food Chem.,
Gatel, F. 1994. Protein quality of legume seeds for non- 50: 197–201.
ruminant animals: a literature review. Anim. Feed Sci. Khokhar, S. and Chauhan, B.M. 1986. Antinutritional factors
Technol., 45: 317–348. in moth bean: varietal differences and effects of methods of
Gemede, H.F. and Ratta, N. 2014. Antinutritional factors in domestic processing and cooking. J. Food Sci., 51: 591–594.
plant foods: potential health benefits and adverse effects. King, T.P., Begbie, R. and Cadenhead, A. 1983. Nutritional
Glob. Adv. Res. Food Sci. Technol., 3(4):103–17. toxicity of raw kidney beans in pigs. Immunocytochemical
Gholami, A., De Geyter, N., Pollier, J., Goormachtig, S., and cytopathological studies on the gut and the pancreas.
and Goossens, A. 2014. Natural product biosynthesis in J. Sci. Food Agric., 34: 1404–1412.
Medicago species. Natural Product Reports, 31: 356-380. Liebman, M. and Okombo, J. 2009. Oxalate content of selected
Hang, D.T., Vanhanen, L. and Savage, G. 2012. Effect of simple pasta products. Journal of Food Composition and Analysis, 22:
processing methods on oxalate content of taro petioles and 254-256.
leaves grown in central Viet Nam. Food Science and Technol., Liener, I.E. 1994. Implications of antinutritional components
pp. 1-5. in soybean foods. Crit. Rev. Food Sci. Nutr., 34: 31–67.
Hegarty, M.P. 1978. Toxic Amino Acids of plant origin. In: Lim, S.J. and Lee, K.J. 2011. A microbial fermentation of
Effect of poisonous plants on livestock. Keeler, R.F., Van soybean and cottonseed meal increases antioxidant
Kampen, K.R. and James, L.F. (eds.) Academic Press New activity and gossypol detoxification in diets for Nile tilapia,
York, pp. 575-585. Oreochromis niloticus. Journal of the World Aquaculture
Society, 42(4): 494–503.

109
 Joshi et al.

Lis, H. and Sharon, N. 1998. Lectins: carbohydrate-specific Quintana, P.J.E., de Peyster, A., Klatzke, S. and Park, H.J. 2000.
proteins that mediate cellular recognition. Chem. Rev., 98: Gossypol-induced DNA breaks in rat lymphocytes are
637–674. secondary to cytotoxicity. Toxicology Letters, 117(1-2): 85–94.
Lyimo, M., Mugula, J. and Elias, T. 1992. Nutritive composition Rafińska, K., Pomastowski, P., Wrona, O., Górecki, R. and
of broth from selected bean varieties cooked for various Buszewski, B. 2017. Medicago sativa as a source of secondary
periods. J. Sci. Food Agric., 58: 535–539. metabolites for agriculture and pharmaceutical industry.
Phytochemistry Letters, 20: 520-539.
Mali, J.M., Kute L.S., Jambhale N.D. and Kadam S.S. 1990.
Effect of leaf processing on antinutrients in leucaena seeds. Ratanapo, S., Ngamjunyaporn, W. and Chulavatnatol, M.
Indian J. Anim. Sci., 60: 385-388. 1998. Sialic acid binding lectins from leaf of mulberry
(Morus alba). Plant Sci., 139: 141–148.
Milgate, J. and Roberts, D.C.K. 1995. The nutritional &
biological significance of saponins. Nutr. Res., 15: 1223– Rogers G. M., Poore M. H. and Paschal J. C. 2002. Feeding
1249. cotton products to cattle. Veterinary Clinics of North America:
Food Animal Practice, 18(2): 267–294.
Mosha, T.C., Gaga, H.E., Pace, R.D., Laswai, H.S. and Mtebek.
1995. Effect of blanching on the content of antinutritional Samtiya, M., Aluko, R.E. and Dhewa, T. 2020. Plant food
factors in selected vegetables. Plant Food Hum. Nutr., 47: anti-nutritional factors and their reduction strategies: an
overview. Food Prod Process and Nutr., 2(6).
361-367.
Savage, G.P., Vanhanen, L., Mason, S.M. and Ross, A.B. 2000.
Nahrstedt, A. 1996. Relationships between the defense
Effect of Cooking on the Soluble and Insoluble Oxalate
systems of plants and insects. In: Romeo, S.B. (Ed.), Recent
Content of Some New Zealand Foods. Journal Food
Advances in Phytochemistry. Plenum Press, New York, pp.
Composition and Analysis, 13: 201-206.
217–230.
Shajeela, P.S., Mohan, V.R., Jesudas, L.L. and Tresina, S.P.
Noftsger, S.M., Hopkins, B.A., Diaz, D.E., Brownie, C. and
2011. Nutritional and Antinutritional Evaluation of Wild
Whitlow, L.W. 2000. Effect of whole and expanded-expelled
Yam (Dioscorea Spp.) [Evaluación Del Valor Nutricional
cottonseed on milk yield and blood gossypol. Journal of
Y Factores Antinutricionales De Dioscorea Spp. Silvestre].
Dairy Science, 83(11): 2539–2547. Tropical and Subtropical Agroecosystems, 14: 723-730.
Noonan, S.C. and Savage, G.P. 1999. Oxalate content of foods Sharma, A. and Sehgal, S. 1992. Effect of processing and
and its effect on humans. Asia Pacific J. Clin. Nutr., 8(1): 64- cooking on the antinutritional factors of faba bean (Vicia
74. faba). Food Chem., 43: 383–385.
Oakenfull, D. and Sidhu, G.S. 1989. Saponins. In: Toxicants Shawrang P., Mansouri M.H., Sadeghi A.A. and Ziaie F. 2011.
of Plant Origin, Vol. 11: Glycosides (Cheeke P, ed.), CRC Evaluation and comparison of gamma- and electron beam
Press, Boca Raton, FL, pp. 97–141. irradiation effects on total and free gossypol of cottonseed
Onabolu, A.O., Oluwole, O.S.A., Rosling, H. and Bokanga, meal. Radiation Physics and Chemistry, 80(6): 761–762.
M. 2002. Processing factors affecting the level of residual Sijun, D., Pawlak, A., Poźniak B. et al. 2012. Effects of gossypol
cyanohydrins in gari. J. Sci. Food Agric., 82: 966‐969 acetic acid on cellular and humoral immune response in
Oscarsson, K.V. and Savage, G.P. 2007. Composition and non-immunized and SRBC-immunized mice. Central-
availability of soluble and insoluble oxalates in raw and European Journal of Immunology, 37(1): 11–19.
cooked taro (Colocasia esculenta var. Schott) leaves. Food Singh, A.B., Awasthi, C.P. and Singh, N. 1999. Biochemical
Chemistry, 101: 559-562. composition and nutritive value of promising collections
Petit, P.R., Sauvaire, Y.D., Hillaire-Buys, D.M., Leconte, O.M., of elephant foot yam (Amorphophallus companulatus Roxb.).
Baissac, Y.G., Posin, G.R, et al. 1995. Steroid saponins Veg. Sci., 26(2): 186-187.
from fenugreek seeds: extraction, purification, and Siritunga, D. and Sayre, R.T. 2003. Generation of cyanogen‐
pharmacological investigation on feeding behaviour and free transgenic cassava. Planta, 217: 367‐373.
plasma cholesterol. Steroids, 60: 674–80. Sirohi, S.K., Pandey, N., Goel, N., Singh, B., Mohini, M.,
Prasad, S. and Dhanya, M.S. 2011. Determination and Pandey, P. and Chaudhry, P.P. 2009. Microbial Activity and
detoxification of cyanide content in sorghum for ethanol Ruminal Methanogenesis as Affected by Plant Secondary
production using Saccharomyces cerevisiae strain. J. Metabol. Metabolites in Different Plant Extracts. International Journal
Syst. Biol., 2(1): 10‐14. of Civil and Environmental Engineering, 1(1): 52-58.
Price, K.R., Johnson, I.T. and Fenwick, G.R. 1987. The chemistry Smith, D.L. 1982. Calcium oxalate and carbonate deposits in
and biological significance of saponins in food and feeding plant cells. In: Anghileri, L.J. and Tuffet, Anghileri, A.M.
stuffs. CRC Crit. Rev. Food. Sci. Nutr., 26: 27–135. Ed. The role of calcium in Biol Systems, CRS Press: Boca
Raton. Florida, pp. 253-262.

110
 Anti-Nutritional Factors in Food and Plant Crops

Soetan, K.O. and Oyewole, O.E. 2009. The need for adequate Van, Damme, E.J.M., Hao, Q., Charels, D., Barre, A., Rouge´, P.,
processing to reduce the antinutritional factors in plants Van Leuven, F. and Peumans, W.J., 2000. Characterization
used as human foods and animal feeds: A review. African and molecular cloning of two different type 2 ribosome-
Journal of Food Science, 3(9): 223-232. inactivating proteins from the monocotyledonous plant
Soetan, K.O. and Oyewole, O.E. 2009. The need for adequate Polygonatum multiflorum. Eur. J. Biochem., 267: 2746–2759.
processing to reduce the anti‐nutritional factors in plants Vetter, J., 2000. Plant cyanogenic glycosides. Toxicon., 38: 11–36.
used as human foods and animal feeds: a review. Afr. J. Vidal, V.C., Frias, J., Estrella, I., Gorospe, M.J., Ruiz, R. and
Food Sci., 3(9): 223–32. Bacon, J. 1994. Effect of Processing on Some Antinutritional
Soto-Blanco, B. 2008. Gossipol e fatores antinutricionais da Factors of Lentils. J. Agric. Food Chem., 42(10): 2291–2295.
soja, In: Toxicologia Aplicada a MedicIna VeterIn aria, H.S. Yalcin, S. and Basman, A. 2015. Effects of infrared treatment
Spinosa, S.L. Gorniak, and J.P. Neto, Eds., Manole, Barueri, on urease, trypsin inhibitor and lipoxygenase activities of
´Brazil, pp. 531–545. soybean samples. Food Chem., 169: 203–10.
Sparg, S.G., Light, M.E. and van Staden, J. 2004. Biological Yildirim, M., Lim, C., Wan, P.J. and Klesius, P.H. 2003. Growth
activities and distribution of plant saponins. J. Ethno- performance and immune response of channel catfish
pharmacol, 94(2–3): 219–43. (Ictalurus puctatus) fed diets containing graded levels of
Tuncel, G., Nout, M.J.R. and Brimer, L. 1995. The effects of gossypol-acetic acid. Aquaculture, 219(1–4): 751–768.
grinding, soaking and cooking on the degradation of Zagrobelny, M., Bak, S. and Møller, B.L. 2008. Cyanogenesis in
amygdalin of bitter apricot seeds. Food Chem., 53: 447‐451. plants and arthropods. Phytochemistry, 69(7): 1457-68.
Uematsu, Y., Hirata, K. and Saito, K. 2000. Spectrophotometric
determination of saponin in Yucca extract used as food
additive. J. AOAC Int., 83: 1451–4.

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