Field Crops Research 115 (2010) 329–339

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Field Crops Research

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Review

Nutritional value of faba bean (Vicia faba L.) seeds for feed and food
Katell Crépon a, Pascal Marget b, Corinne Peyronnet a, Benoit Carrouée a, Paolo Arese c, Gérard Duc b,*
a
Union nationale interprofessionnelle des plantes riches en protéines (UNIP), 12 av. George V, 75008 Paris, France
b
Institut National de la Recherche Agronomique (INRA), UMR 102, Génétique et Ecophysiologie des Légumineuses à Graines, BP 86510, 21065 Dijon cédex, France
c
Dipartmento di Genetica, Biologia e Biochimica, Università di Torino, Via Santena 5 bis, 10126 Torino, Italy

A R T I C L E I N F O A B S T R A C T

Article history: Faba bean (Vicia faba L.) is a protein-rich legume seed well adapted to most climatic areas of Europe and
Received 19 September 2009 widely used for feed and food. Even if the seed is generally recognized to be of good nutritional value, existing
Received in revised form 24 September 2009 genetic variability for seed composition offers possibilities for improvement of this trait by breeding. Four
Accepted 24 September 2009
major quality types must be distinguished according to the presence or absence of tannins in the
integuments and of vicine (V) and convicine (C) in the cotyledons. The nutritional value of diets containing
Keywords: varying amounts of different faba bean cultivars characterized by high or low levels of tannins, and high or
Faba bean
low levels of vicine + convicine (VC), has been examined in monogastric animals and ruminants. Low-tannin
Vicia faba L.
Tannins
content generally results in higher protein and energy digestibility for monogastric animals and low VC
Vicine/convicine content has a positive effect on laying hen and broiler production performances. V and C, inactive precursors
Animal nutrition of divicine and isouramil are redox compounds potentially toxic to human carriers of a widespread genetic
Human nutrition deficiency of the erythrocyte (red blood cell, RBC) enzyme glucose-6-phosphate dehydrogenase (G6PD).
Favism Ingestion of faba beans by these deficient individuals may cause a severe, potentially lethal hemolytic
Technological treatments anemia (favism). The mechanism of action of divicine and isouramil in the G6PD-deficient RBC is discussed.
Beside the positive impact of using tannin-free varieties in monogastric animals diets, the
development of faba bean cultivars with very low levels of VC would represent a real advantage in terms
of nutritional performance in poultry diets and of food safety to humans.
ß 2009 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 330
2. Genetics of seed composition in faba beans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 330
3. Faba beans in animal nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331
3.1. Nutritional values . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331
3.1.1. Nutritional values for pigs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331
3.1.2. Nutritional values for poultry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
3.1.3. Nutritional value for ruminants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
3.2. Faba bean in diets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334
3.2.1. Faba bean in pig diets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334
3.2.2. Faba bean in poultry diets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
3.2.3. Faba bean in ruminants diets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
4. Faba beans in human nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
4.1. Potential toxicity of faba beans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
4.2. Favism in G6PD-deficient human subjects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
4.3. Faba beans, malaria and G6PD deficiency . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
5. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 338
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 338

Abbreviations: V, vicine; D, divicine; C, convicine; I, isouramil; VC, vicine + convicine; T+, tannin-containing; T , tannin-free; RBC, red blood cell, erythrocyte; GSH, reduced
glutathione; G6PD, glucose-6-phosphate dehydrogenase; LDL, low-density lipoprotein.
* Corresponding author. Tel.: +33 3 80 69 31 48; fax: +33 3 80 69 32 63.
E-mail addresses: marget@dijon.inra.fr (P. Marget), b.carrouee@prolea.com (B. Carrouée), paolo.arese@unito.it (P. Arese), Gerard.Duc@dijon.inra.fr, duc@dijon.inra.fr
(G. Duc).

0378-4290/$ – see front matter ß 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.fcr.2009.09.016
330 K. Crépon et al. / Field Crops Research 115 (2010) 329–339

1. Introduction Table 1
Chemical composition of faba bean seeds (dry matter basis) of varieties with low- or
high-tannin content.
With their seeds rich in protein and energy and their ability to
grow in various climatic zones, faba bean (Vicia faba L.) production Duc et al. (1999) Sauvant et al. (2004)
Meana Mean (SD)
has a long history of numerous and valuable uses in feed and food.
Nevertheless, faba bean seeds contain different constituents that High-tannin faba beans
may exert anti-nutritional effects. As far as animal nutrition is Crude protein (g/kg) 310 294 (25)
Starch (g/kg) 412 443 (31)
concerned, tannins, vicine (V), and convicine (C) are faba bean seed
Crude fibre (g/kg) 99 91 (13)
constituents which have been demonstrated in several studies to Sugars (g/kg) 38 35 (9)
have an anti-nutritional effect in the diet of monogastric animals Fat (g/kg) 19 15 (4)
(Olaboro et al., 1981; Grosjean et al., 2000). In human nutrition, it TIA (UTI/mg)b 2.9
Condensed tannins (g/kg) 6.6
was demonstrated that divicine and isouramil, active derivatives of
Vicine + convicine (g/kg) 8.3
V and C contained in the seed, are toxic to human carriers of a Lysine (g/kg) 20.3 19.2
widespread genetic defect (deficiency of the erythrocyte-located Methionine (g/kg) 2.7 2.1
glucose-6-phosphate dehydrogenase (G6PD)). After a 30-year Cysteine (g/kg) 3.9 3.7
period of breeding efforts in Europe (1970–2000), a large genetic Tryptophane (g/kg) 2.7 2.4

variability was shown to exist within the V. faba species, which has Low-tannin faba beans
led to improvements of yield potential and seed composition (Bond Crude protein g/kg) 319 311 (26)
and Duc, 1993; Duc, 1997). Currently, in the European catalogue of Starch (g/kg) 427 433 (27)
Crude fibre (g/kg) 88 87 (10)
registered varieties, four types of faba beans can be distinguished Sugars (g/kg) 44 43 (8)
according to the content of tannins in the integuments (low or Fat (g/kg) 20 13 (2)
high) and the content of vicine + convicine (VC) in the cotyledons TIA (UTI/mg)b 2.9
(low or high) (Duc et al., 1999). However, the most frequently Condensed tannins (g/kg) 0.1
Vicine + convicine (g/kg) 7.6
cultivated types still have high-tannin and VC contents and a
Lysine (g/kg) 19.5 20
potential of improvement of their nutritional value through Methionine (g/kg) 2.6 2.2
breeding exists. Cysteine (g/kg) 3.6 3.9
Tryptophane (g/kg) 2.7 2.6
2. Genetics of seed composition in faba beans a
Means of four low-tannin lines carrying gene zt1, compared to the mean of their
four high-tannin isogenics.
b
Single seed weight varies from 0.2 up to 2.6 g of DM in faba bean UTI: units of trypsin inhibitor activity (Valdebouze et al., 1980).
(V. faba L.) genetic resources. Two types of varieties are cultivated:
(1) cultivars with large flattened seeds, from 1 to 2 g DM per seed, (Duc et al., 1999) and faba bean commercial varieties (Sauvant
named ‘‘V. faba major or broad beans’’, mainly grown in the et al., 2004) are compared.
southern regions of Europe and used for human food, either as In Europe, most of the T+ cultivars presently grown have a mean
fresh seeds or as dry seeds; (2) varieties with medium to relatively seed tannin content of 5–10 g/kg DM (Duc et al., 1999) and in
small and round seeds, from 0.4 to 0.8 g DM per seed, named ‘‘V. relation with this trait, their flowers display a large black spot on
faba minor or field beans or horse beans’’), grown in a larger range the wings, their standard being often coloured in pale pink, pink or
of regions, mainly for dry seeds used for animal feed or for human red. Throughout this paper we use the generic term ‘‘tannins’’ to
food. In small seeds, the DM proportion of integument is higher mean proanthocyanidins, i.e. condensed tannins. These polyphe-
than for large seeds. Duc et al. (1999) found values from 137 to nolic compounds are concentrated in the seed coats. Mutant alleles
146 g/kg of hulls in seed dry matter (DM) for tannin-containing at two distinct genetic loci (zt1 and zt2) involved in tannin
(T+) varieties, higher than for tannin-free (T ) varieties which synthesis have been discovered in V. faba genetic resources (Picard,
varied from 110 to 134 g/kg DM. In relation with this tannin trait, 1976), either of which reduces the seed tannin content and
these authors found a mean wall insoluble cell wall (WICW) simultaneously determines a white flower colour. A few T
content in the seed of 184 g/kg seed DM in T+ cultivars (61% cultivars with white flowers have been released in Europe with a
contained in seed coats) higher than for T cultivars which mean condensed tannin content of 0.1 g/kg DM (Bond and Duc,
contained 167 g/kg of WICW (57% contained in seed coats). Protein 1993). Gene zt2 may offer a nutritional advantage over zt1 by also
content in faba bean seeds ranges from 247 to 372 g/kg DM in increasing seed protein level and reducing fibre content (Duc et al.,
genetic resources (Duc et al., 1999) and from 270 to 320 g/kg DM in 1999; Grosjean et al., 2001). Seeds of T varieties have a
commercial varieties. This trait usually shows high heritability significantly higher in vitro digestibility of the dry matter, on
(Link et al., 2005) and can be efficiently selected. Even if some average 4–7% higher than T+ varieties (Bond, 1976). The fact that
genetic variability has been measured for the trypsin inhibiting these two recessive alleles at distinct loci are available to breed for
activity of the proteins, this is generally very low, below 5 units/mg the T lines implies careful isolation of varieties during the
DM and does not appear as a limiting factor in animal nutrition multiplication process of this allogamous species. Indeed uncon-
(Valdebouze et al., 1980; Bond and Duc, 1993; Duc et al., 1999). trolled hybridisation between zt1 and zt2 or with wild types lines
Starch is the major faba bean seed component; it has a mean would produce T+ progenies. The double recessive genotype zt1–
content of 423 g/kg DM and varies in negative correlation with zt2 has never been obtained which suggests a possible lethality for
protein content (Duc et al., 1999). A detailed analysis on 12 such genotypes (Crofton and Bond, 1998; Crofton et al., 2000).
genotypes revealed low fat and sugar contents in the seeds (19 and Recently molecular markers linked to T character were devel-
41 g/kg DM, respectively) (Duc et al., 1999). In comparison with oped which offer a possibility of early breeding with a marker-
cereal, faba bean seeds have high lysine content (19.8 g/kg DM) assisted strategy for this trait (Gutierrez et al., 2008). There is a
and low methionine, cysteine and tryptophan content (2.6, 3.7 and relationship between seed coat colours and tannin content in the
2.7 g/kg DM) (Mossé, 1990; Duc et al., 1999). In practice, mixtures seeds. According to genotypes, seed coat colours can be clear-
of faba beans with other protein sources in diets allow to meet beige, grey-beige, grey-green, grey or dark-grey for T lines and
animal requirements. Composition data for coloured and white black, brown, red, violet, green, yellow or beige for T+ lines. When
faba beans are compiled in Table 1 where isogenic pairs (T+ vs. T ) aging, the seed colour remains stable in T lines, whereas it
 K. Crépon et al. / Field Crops Research 115 (2010) 329–339 331

becomes darker in T+ lines. Therefore, T+ vs. T cultivar distinction
is not always accurate on the basis of seed colours, because of the
aging effect and the overlap in colours between the two groups
(Crofton and Bond, 1998).
V and C are glucopyranosides concentrated in cotyledons of
faba bean seeds (Olaboro et al., 1981; Griffiths and Ramsay, 1993).
In conventional cultivars, VC content ranges from 6 to 14 g/kg of
mature seed DM (Duc et al., 1999). V and C are thermostable seed
constituents not easily removed by technological processes. In V.
faba genetic resources, a mutant allele vc has been discovered
which reduces 10–20-fold V and C contents (Duc et al., 1989,
1999). The effect of vc in relation with wild allele is additive, and
the VC content in the seed is determined by the mother plant
genotype (Duc et al., 1989). In breeding programs, measurements
of VC is performed by expensive HPLC measurements or rapid
colorimetric test, but the hilum colour can be used as phenotypic
Table 2
Nutritional values for pigs of faba beans varieties with low- or high-tannin content, low- or high-(vicine + convicine) (VC) content.
marker, the two genes involved in these traits being calculated as
10.1 centimorgans apart (Duc et al., 2004a,b). Molecular markers
were also developed to assist the selection for low VC content
genotypes (Gutierrez et al., 2006). In 2008, six low VC content
varieties were present in the French catalogue. A very new variety
type combining T and low VC characters, called FEVITA1 (Duc
et al., 2004a,b) also recently started to be bred: the first variety of
this type has been registered in the French catalogue in 2004.
3. Faba beans in animal nutrition
3.1. Nutritional values
3.1.1. Nutritional values for pigs
The impact of V and C on the nutrient digestibility in pigs is not
very well documented but it seems to be negligible (Grosjean et al.,

Cultivar Apparent digestibility Digestible energy Ileal apparent digestibility Reference

content (MJ/kg DM)
Crude protein (%) Energy (%) Crude protein (%) Lysine (%)

High-tannin faba beans (T+)

High VC content
Talo 71.9 78.5 14.5 Bourdon and Perez (1984)
Alfred 81.8 86.7 Maillard et al. (1990)

Mythos 82.4 82.1a 15.3b 72.4 82.1 Jansman et al. (1993)

Alfred 79.4 79.0a 14.7b 71.5 84.3

Alfred 80.5 15.1 82.7 79.8 Grosjean et al. (2001)

Castel 80.5 15.0 84.2 88.5

NR1 82.9 76.7 14.4 Grosjean et al. (2001)

Fabiola 78.9 78.5 14.5
NR2 79.2 77.9 14.7
Robin 79.3 79.6 14.8

Maya 82.0 82.0 15.2 82.5 90.7 Vilariño et al. (2004)

Méli 76.7 85.3
Olan 77.6 87.4

Low VC content
NR3 79.5 81.0 15.2 Grosjean et al. (2001)
Divine 80.3 79.3 14.9 82.3 88.3 Vilariño et al. (2004)

Mean T+ 79.6 79.6 14.9 81.1 85.9

Min 71.9 76.7 14.4 76.7 79.8
Max 82.9 82.0 15.2 84.2 90.7

Low-tannin faba beans (T )

High VC content
NR4 80.2 80.3 14.8 Bourdon and Perez (1984)
NR5 75.8 80.9 15.1

Blandine 84.6 88.2 Maillard et al. (1990)

Blandine 89.3 88.1a 16.4b 85.7 91.6 Jansman et al. (1993)
Albatross 90.3 17.1 Grosjean et al. (2001)

NR6 90.7 94.3 17.5 Grosjean et al. (2001)

Fabiola 85.2 86.3 15.8
NR7 86.7 89.8 16.7

Gloria 73.9 83.3 15.4 78.8 88.3 Vilariño et al. (2004)

Low VC content
NR8 77.9 85.1 Hess et al. (2000)
NR9 78.4 84.0
NR10 75.8 85.8

NR11 89.1 88.2 16.4 Grosjean et al. (2001)

NR12 90.2 91.6 17.1

NR13 83.2 89.2 Vilariño et al. (2004)

Mean T 84.6 87.3 16.2 80.6 87.5

Min 73.9 80.3 14.8 75.8 84.0
Max 90.7 94.3 17.5 85.7 91.6

NR: different genotypes, not-registered varieties.

a
Apparent faecal digestibility of organic matter.
b
Digestible energy calculated from organic matter digestibility.
332 K. Crépon et al. / Field Crops Research 115 (2010) 329–339

Table 3
Nutritional values for poultry of faba beans varieties with low- or high-tannin (T) content, low- or high-vicine and convicine (VC) content.

Cultivar Mash diet Pelleted diet Reference

Apparent digestibility AMEn Apparent digestibility AMEn

(MJ/kg DM) (MJ/kg DM)
Starch (%) Protein (%) Starch % Protein %

High-tannin faba beans (T+)

Adult cockerels
High VC content (T + VC+)
Alfred 12.05 Grosjean et al. (1995)
Victor 12.48
Castel

NR14 95.2 74.0 11.68 97.2 74.7 12.07 Grosjean et al. (2000)
Fabiola 91.7 71.3 10.63 78.5 11.57
NR15 92.3 84.3 11.65 81.6 12.01
Robin 91.8 71.0 11.44
Bourdon 90.5 68.4 10.71

Maya 11.92 Brévault et al. (2003)

Marcel 12.81 (2) Métayer and Vilariño (2006)

Mean T + VC+ for adult 92.3 73.8 11.2 97.2 78.3 12.2

Low VC content (T + VC )
NR16 92.6 84.4 11.67 Grosjean et al. (2000)
Divine 12.76 Brévault et al. (2001)
Divine 11.77 Métayer et al. (2003)

Mean T + VC for adult 92.6 84.4 11.7 12.3

Broiler chicken
High VC content (T + VC+)
Alfred 85.6 66.9 11.16 91.5 (2) 70.6 (2) 11.53 (2) Lacassagne et al. (1988)
Soravi 83.4 69.4 10.82 90.9 (2) 72.2 (2) 11.55 (2)

Alfreda 90.2 69.8 9.55 Lacassagne et al. (1991)

Alfredb 70.3 69.4 12.38

Marcel 12.83 (2) Métayer and Vilariño (2006)

Mean T + VC+ for broiler 82.4 68.9 11.0 91.2 71.4 12.0

Low VC content
Divine 12.31 Métayer et al. (2003)

Mean T + VC for broiler 12.3

Low-tannin faba beans (T )

Adult cockerels
High VC content (T VC+)
Albatross 12.01 Grosjean et al. (1995)
Caspar 12.32

NR17 95.8 83.9 11.83 Grosjean et al. (2000)

Fabiola 92.6 84.4 11.44 87.7 12.45
NR18 92.3 84.3 12.21 87.2 12.46
Glacier 91.8 83 11.25

Gloria 13.22 Brévault et al. (2001)

Gloria 13.38 Brévault et al. (2003)
Gloria 12.10 Métayer et al. (2003)

Gloria 13.0 (2) Métayer and Vilariño (2006)

Victoria 13.24 (2)

Mean T VC+ for adult 93.1 83.9 11.7 87.5 12.8

Low VC content (T VC )
NR19 93.3 81.7 12.21 Grosjean et al. (2000)
NR20 95.7 84.4 12.77 98.8 87.9 13.45

NR21 12.56 Métayer et al. (2003)

Disco 13.08 (2) Métayer and Vilariño (2006)

Mean T VC for adult 94.5 83.1 12.5 98.8 87.9 13.0

Broiler chicken
High VC content (T VC+)
Blandine 75.1 82.6 10.17 86.0 (2) 86.8 (2) 11.9 Lacassagne et al. (1988)

Blandinea 80.4 82.5 11.29 Lacassagne et al. (1991)

Blandineb 63.8 84.2 9.2

Gloria 11.93 Métayer et al. (2003)

 K. Crépon et al. / Field Crops Research 115 (2010) 329–339
Table 3 (Continued )
Cultivar Mash diet
Apparent digestibility AMEn
(MJ/kg DM)
Starch (%) Protein (%)
Gloria
Victoria
Mean T VC+ for broiler 73.1 83.1 10.2
Low VC content (T VC )
NR22
Disco
Mean T VC for broiler
NR: different genotypes, not-registered varieties and (number of measures).
a
Fine grinding.
b
Coarse grinding.
2001; Vilariño et al., 2004). In contrast (Jansman et al., 1995), in an
experiment including high proportions of integuments with or
without tannins, conclude that tannins reduce the true digest-
ibility, increase the endogenous excretion of proteins, and that
they seem to interact preferably with proteins with a high content
of proline and histidine. Several other results, comparing T+ and T
seeds, observed a negative impact of tannin on apparent crude
protein digestibility (Table 2). Concerning ileal apparent digest-
ibility of protein and lysin, which is supposed to be more adapted
to balance pig diets, results on whole seeds are contradictory:
Jansman et al. (1993) observed a strong effect of tannins (91.6% for
lysine in T seeds vs. 83% in T+ seeds) whereas Maillard et al.
(1990) and Vilariño et al. (2004) found no or very little difference.
Concerning energy apparent digestibility and digestible energy,
all authors found lower values for T+ than for T faba beans, yet
with noticeable differences both for the absolute values and for the
difference between T+ and T seeds. On average, however, the
effect of tannins seems to be rather important: the mean digestible
energy of the 10 references with T seeds is 16.2 MJ/kg DM, vs.
14.9 for the 12 references with T+ seeds quoted in Table 2. It can be
noticed that the references published in INRA-AFZ (tables of
composition and nutritional values of a wide range of raw
materials by Sauvant et al., 2004) give a rather higher energy
value for T+ faba bean (15.6 MJ/kg DM) and a similar value for T
faba beans (16.1 MJ/kg DM).
It can be concluded that tannins in faba beans reduce their
nutritional value for pigs, both for energy and for protein, but it is
difficult to quantify precisely the tannin effect, due to the large
variability of the references. This variability is probably due to
interactions with other seed components, such as insoluble cell
walls, and to the variability of the tanning power of the tannins. In
order to answer more precisely this question, experiments with
reconstituted faba beans by mixing T+ or T integuments with
different proportions of cotyledons, would be probably more
conclusive.
3.1.2. Nutritional values for poultry
Because VC and tannins have an impact on the nutritional value
of faba beans for poultry, it is necessary to distinguish all the high
vs. low combinations, which implies four types of faba bean
(Table 3).
Tannins have a significant negative impact on protein digest-
ibility, as shown by Grosjean et al. (2000), Wareham et al. (1993),
Lacassagne et al. (1991), Vilariño et al. (2009). Tannins also reduce
energy (Grosjean et al., 2000; Vilariño et al., 2009) and starch
(Lacassagne et al., 1988) digestibilities.
For high VC and T+ faba beans fed in mash diets to adult
cockerels, the AMEn (apparent metabolisable energy) values
333
Pelleted diet Reference
Apparent digestibility AMEn
(MJ/kg DM)
Starch % Protein %
12.99 Métayer and Vilariño (2006)
12.84
86 86.8 12.4
12.7 Métayer et al. (2003)
12.89 Métayer and Vilariño (2006)
12.8
average 11.22 MJ/kg DM, compared with 11.67 MJ/kg DM for the
low VC and T+ faba bean. In the laying hen, Olaboro et al. (1981)
demonstrated that VC reduced egg size and Muduuli et al. (1981)
related this decrease to erythrocyte hemolysis. For the high VC
and T faba beans fed to adult cockerels in mash diets, the AMEn
values average 11.68 MJ/kg DM compared with 12.49 MJ/kg DM
for low VC and T faba beans. AMEn values in broiler chicken are
less well documented but, as for peas, AMEn values in broiler
chicken are lower than in adult cockerels. The lowering of VC and
tannin contents in faba bean seeds has a significant and additive
positive impact on AMEn values in broiler chickens (Vilariño et al.,
2009).
Pelleting has a very positive and significant effect on the AMEn
value of faba bean fed to young chicks (Lacassagne et al., 1988). The
increase in the AMEn value that can be attributed to pelleting is
about 1.23 MJ/kg DM (12% of the AMEn value of unpelleted faba
bean). The same effect can be observed in other grain legumes such
as pea, the beneficial effect of pelleting being explained by a better
starch and protein digestibility (Carré et al., 1987). In pelleted diets,
high VC and T+ faba beans have mean AMEn values of 12.19 MJ/kg
DM compared with 12.27 MJ/kg DM for low VC and T+ faba beans.
For high VC and T faba beans, mean AMEn values are 12.77 MJ/kg
DM compared with 13.04 MJ/kg DM for low VC and T faba beans.
3.1.3. Nutritional value for ruminants
As for the other grain legume raw seeds, the nitrogen fraction of
faba bean is very soluble and easily degradable in the rumen. With
the ‘‘in sacco’’ method, Faurie et al. (1992) showed that more than
85% of the nitrogen is degraded in the rumen in 2 h resulting in a
mean theoretical nitrogen degradability (DT) of 92% for three
different cultivars. These data are higher than the values reported
by Sauvant et al. (2004) of 83 and 82%, respectively for T and T+
faba beans.
Technological treatments may have an impact on the
degradability of nitrogen of faba bean. Nitrogen degradability
decreases when the grinding screen changes from 0.8 to 3.0 mm
by about 10%, whatever the cultivar (T+ or T lines), as shown
by Foucher et al. (unpublished). A finer grinding results in a
higher nitrogen degradability and consequently in a lower value
of digestible protein in the intestine (PDI value) (Table 4).
Extrusion may protect nitrogen from degradability in the rumen,
as shown by Cros et al. (1991). On the contrary, the presence of
tannin in the hull does not seem to protect nitrogen from
degradability, considering the results obtained by Foucher et al.
(unpublished).
The energy values of faba bean for ruminant (UFL and UFV) are
high, comparable to cereal energy values (Sauvant et al., 2004) and
explained by the high starch content of the seeds.
334 K. Crépon et al. / Field Crops Research 115 (2010) 329–339

Table 4
Nutritional values for cows of faba beans varieties with low- or high-tannin content.

Cultivar Treatment UFL UFV PDIA PDIN PDIE DT dr dMO Reference

(kg DM) (kg DM (kg DM) (kg DM) (kg DM) (%) (%) (%)

Talo (HT) Raw 89.2 Cros et al. (1991)

Extruded 120 8C 70.2

Gloria (LT) Grinding 0.8 mm 1.37 1.41 43 205 127 86.6 99.3 Foucher et al.
Grinding 3 mm 1.28 1.28 68 211 144 77.8 93.9 (2002, unpublished)

INRA00 (LT) Grinding 0.8 mm 1.35 1.37 35 176 118 86.6 97.9 Foucher et al.
Grinding 3 mm 1.29 1.32 57 183 134 78.6 95.7 (2002, unpublished)

Divine (HT) Grinding 0.8 mm 1.25 1.25 43 205 127 83.3 92.6 Foucher et al.
Grinding 3 mm 1.17 1.16 67 188 138 75.1 88.2 (2002, unpublished)

INRA 2004 (LT) 1.19 1.19 52 197 113 83.0 89.0 91.0 Sauvant et al. (2004)
INRA 2004 (HT) 1.2 1.2 52 187 112 82.0 89.0 91.0

LT: low-tannin; HT: high-tannin; UFL: energy value for milk; UFV: energy value for meat; PDIA: protein digestible in the intestine from dietary origin; PDIN:
PDIA + (PDIMN = PDI from microbial origin depending on degradable nitrogen); PDIE: PDIA + (PDIME = PDI from microbial origin depending on fermentescible energy); DT:
theoretical degradability of N; dr: real degradability of N; dMO: degradability of organic matter.

3.2. Faba bean in diets
3.2.1. Faba bean in pig diets
Contrasting results were obtained in former trials (Table 5)
with piglets fed T and T+ faba beans. The trials described by
Fekete et al. (1985) led to the recommendation that inclusion
levels of faba beans should be limited to 100 g/kg in piglet diets
whatever the type of faba bean variety. However, recent trials
showed that feeding piglets with faba beans at 200 g/kg, with or
without tannins, would lead to performances similar to the
control diets (Euronutrition, 2003, unpublished; Royer et al.,
2009, submitted). This may be due to a better amino acid
balance in the diets compared to older trials which were based
on maize.

Table 5
Performance of piglets fed faba bean-containing diets, made with low- or high-tannin varieties.

Diet Faba bean Amino acid Weight at the Average daily Feed convertion Tannin Reference
(g/kg) supplement beginning (kg) gain, % (g/day) ratio, % (g/day) variety type

Wheat/barley/maize 0 10.8 100 100 Fekete et al. (1985)

100 101 102 Low-tannin
200 92** 108** Low-tannin
300 85** 114** High-tannin

Wheat/barley/maize 0 8.4 100 100 Fekete et al. (1985)

100 110** 98 Low-tannin
200 104 99 Low-tannin
300 96 112** Low-tannin

Wheat/barley/maize 0 9.15 100 100 Fekete et al. (1985)

100 100 107 Low-tannin
200 99 102 Low-tannin
300 85 104 Low-tannin

Wheat/barley/maize 0 10.9 100 100 Fekete et al. (1985)

100 98 102 High-tannin
150 96 103** High-tannin
200 93** 105** High-tannin

Wheat/barley/maize 0 10.1 100 100 Fekete et al. (1985)

100 100 102 High-tannin
150 95 103** High-tannin
200 93** 105** High-tannin

Wheat/barley/maize 0 10.9 100 100 Fekete et al. (1985)

150 98 104 Low-tannin
150 100 104 High-tannin
150 97 105 High-tannin

Triticale/barley/maize 0 8.0 100 (511) 100 (1.82) Skiba (2000)

80 90 102.2 High-tannin
150 96 100 High-tannin
200 91.4 101.6 High-tannin
0 12.3 100 (578) 100 (1.71) Euronutrition
100 99.6 101.2 Low-tannin (2003, unpublished)
200 100.3 97.1 Low-tannin

Wheat/barley 0 Lysine HCl, 8.0 100 (544) 100 (1.52) Royer et al.
100 methionine, 100.2 98.0 Low-tannin (2009, submitted)
100 threonine, 97.6 100.6 High-tannin
200 tryptophane 100.7 100.6 Low-tannin
200 99.6 101.3 High-tannin
**
P < 0.01.
 K. Crépon et al. / Field Crops Research 115 (2010) 329–339 335

Table 6
Performance of growing and finishing pigs fed faba bean-containing diets made with low- or high-tannin varieties.

Diet Faba Amino acid Average daily Feed Dressing Lean Backfat, Tannin Reference
bean supplement gain, % (g/day) convertion percentage percentage % (mm) variety
(g/kg) ratio (%)a (%) (%) type

Maize 0 100 (692) 100 (3.08) 100 (74.3) Bourdon and

150 98.8 99.0 99.9 High-tannin Perez (1976)
300 99.3 102.9 99.3 High-tannin

Barley 0 100 (525) 100 (3.76) Henry and

150 106.3 94.4 High-tannin Bourdon
330 95.0 103.2 High-tannin (1978)
330 Methionine 95.6 103.7 High-tannin

Maize 0 100 (567) 100 (3.39) Henry and

330 82.5 107.9 High-tannin Bourdon
330 Methionine 75.8 113.6 High-tannin (1978)

Maize 0 100 (616) 100 (3.12) Henry and

150 97.4 104.5 High-tannin Bourdon
150 Tryptophane 100.3 100.6 High-tannin (1978)
300 91.9 110.6 High-tannin
300 Tryptophane 98.2 103.2 High-tannin

Maize 0 100 (761) 100 (2.92) 100 (77.4) 100 (50.6) 100 (22.5) ITCF (1983)
175 94.9 105.8 99.7 100.6 97.3 Low-tannin
175 97.1 103.1 99.1 101.6 96 High-tannin
175 94.7 105.5 99.5 99.8 96 High-tannin

Maize 0 100 (768) a 100 (2.88) a 100 (77.5) 100 (49.7) 100 (23.1) ITCF-AGPM
100 100.4 a 100.3 a 100.1 98.4 104.3 High-tannin (1985)
150 94.5 b 106.2 b 99.7 98.9 100 High-tannin
200 93.3 b 107.9 b 99.7 101.2 96.1 High-tannin

Maize 0 100 (761) 100 (2.89) 100 (76.8) 100 (49.6) 100 (24.2) ITCF-AGPM
220 98.9 102.4 99.9 104 91.7 High-tannin (1986)
220 Methionine 98.2 103.5 99.9 102.4 91.7 High-tannin
(0.04%)
220 Methionine 96.6 104.8 99.7 101 96.3 High-tannin
(0.08%)

0 100 (680) 100 (2.75) Thacker and

100 97 106.2 High-tannin Bowland
150 95.5 105.0 High-tannin (1985)
200 95.5 106.9 High-tannin
250 89.7 118.5 High-tannin
300 85.3 123.6 High-tannin

0 100 (988) 100 (2.71) 100 (74.8) 100 (59.2) Euronutrition

150 97.8 101.5 100.5 100.3 Low-tannin (2003,
300 100.2 98.1 100.0 101.5 Low-tannin unpublished)

Wheat/ 0 Methionine + 100 (962) 100 (2.76) 100 (60.3) Royer et al.
barley/ 200 threonine + 100.1 100.0 98.8 Low-tannin (2009,
maize 350 tryptophane 97.2 102.9 99.2 Low-tannin submitted)
200 96.2 103.9 99.8 High-tannin
350 98.7 101.4 99.3 High-tannin

Letters: groups of significant differences P < 0.05.

a
Ratio feed consumption/weight gain.

Previous trials carried out with T or T+ faba beans in growing
or finishing pig diets (Table 6) have shown that the use of faba
beans at up to 100 g/kg did not alter growth performance. At rates
in excess of 100 g/kg, the inclusion of faba beans tended to reduce
the animal growth and to increase the feed conversion ratio,
whatever the tannin type of faba beans. Supplementation with
tryptophane tends to reduce the difference in performance
compared with the control diet (Henry and Bourdon, 1978),
whereas supplementation with methionine does not have any
effect (ITCF-AGPM, 1986). However, two recent trials (unpub-
lished) show that in diets supplemented with amino acids, the
inclusion of T or T+ faba beans at up to 350 g/kg, does not alter
growth performance. This result may be due to a better amino acid
balance, a formulation of digestible amino acids and a better
knowledge of faba bean energy values.
The introduction of T+ faba bean at a rate of 150 g/kg in the diets
of gestating and lactating sows had no effect on reproductive
performances (number of piglets born and number of litters
weaned) (Buron and Gatel, 1992). In this previous trial, the effect of
faba bean on milk production has not been measured, but
according to Etienne et al. (1975), the inclusion of 150 g/kg of
faba bean in the diet, supplemented with methionine, does not
affect the quantity and quality of milk produced.
3.2.2. Faba bean in poultry diets
Lucbert (1986) reviewed several trials showing that the use
of faba bean in broiler diets leads to a slight increase in the feed
conversion ratio, due to an increase in feed intake and a decrease
in daily growth. Nevertheless, supplementing the diet with
methionine dramatically improves performance (Marquardt and
Fröhlich, 1981). In a more recent trial with T+ and T faba beans
(Table 7), it was shown that the inclusion of both faba bean
types at 250 g/kg resulted in similar performances to that
obtained with the control diet (Métayer et al., 2003). These
336 K. Crépon et al. / Field Crops Research 115 (2010) 329–339
Table 7
Performance of broiler chicken fed faba bean-containing diets made with low- or high-tannin varieties.
Performance of diet Control diet with soya Diet with of faba bean
Gloria (low-tannin) 250 g/kg
Live weight gain (g/d) 52.1 51.1
Food intake 1–56 d (g) 5844 5635
Feed conversion ratio 1–56 d 2 1.97
Gloria (low-tannin) 200 g/kg
Live weight gain (g/d) 31.4 ab 31.8 b
Food intake 1–83 d (g) 7385 b 7341 b
Feed conversion ratio 1–83 d 2.87 2.82
d: day.
P: probability of significant difference.
Letters: groups of significant differences P < 0.05.
results are not in accordance with those obtained by Brévault
et al. (2003) (Table 7), who noticed a significant decrease in
performance with 200 g/kg of T+ faba bean (cultivar Maya)
compared with the control diet, or with the diet including T
faba bean (cultivar Gloria).
In laying hen, it has been suggested that the depressing effect
on egg size by VC (Olaboro et al., 1981) may be related the
production of derivative pro-oxidants that cause lipid peroxida-
tion, hemolysis and interfere with normal lipid metabolism in the
hen (Muduuli et al., 1981). When faba bean is incorporated into
the diets of laying hens at rates up to 7%, a significant decrease in
egg weight is noticed, and this cannot be improved using
tryptophane, methionine or linoleic acid supplements (Lacas-
sagne, 1988). In a recent trial, Lessire et al. (2005) introduced 20%
of two faba bean cultivars, one with a high content of VC (cv.
Marcel) and the other with a low VC content (cv. Divine). The
laying rate was not modified by the two experimental diets, but
egg weight was negatively correlated with dietary VC content. Egg
quality was not impaired by faba bean. The use of faba bean with a
high VC content cannot exceed 7% of the diet, but it is possible to
include faba bean at up to 20% if the cultivar used has a low VC
content.
3.2.3. Faba bean in ruminants diets
Several trials have tested faba bean as a protein source in dairy
cows diets in order to replace soybean meal. Brunschwig and
Lamy (2002) showed that the use of 30% of ground faba bean in the
concentrate feed for dairy cows does not alter the feed
consumption, the milk production (higher than 30 kg per cow
per day) or the milk composition (crude protein or lipid). It
represents an average consumption of faba bean of 3.5 kg per day
and per animal. Nevertheless, a decrease in protein content in milk
can be observed with feed rich in faba bean (4.5 kg per cow per
day) (Trommenschlager et al., 2003). This can result from the
higher level of urea in the milk which was also observed by
Brunschwig et al. (2004), who explain it by the high solubility of
nitrogen in the rumen.
Lapierre et al. (2003) substituted all the soybean meal of the
cow diets by a mixture of 1/3 of rapeseed meal and 2/3 of
laminated faba bean (with 3.7 kg per cow and per day of faba bean).
With iso-nitrogen (PDI) and -energy (UFL) based diets, there were
no differences on milk production and composition. These trials
demonstrate the good value of faba bean as a component of
concentrate for dairy cows receiving well-balanced diets without
any urea supplementation.
The use of faba bean, instead of soybean meal, in concentrate
feed for lambs does not decrease feed consumption, animal growth
and the composition of carcass (Di Grigoli et al., 2005; Delmotte
and Rampanelli, 2006). Nevertheless, these results are subject to
the presentation of feed: faba bean must be presented crushed, and
P Reference
Divine (high-tannin) 250 g/kg Métayer et al. (2003)
51.8 NS
5716 NS
1.98 NS
Maya (high-tannin) 200 g/kg Brévault et al. (2003)
30.5 a <0.01
7135 a <0.001
2.85 NS
not whole, and mixed with the cereals (Delmotte and Rampanelli,
2006).
Concerning Simmental growing bulls, Leitgeb and Lettner
(1992) showed that the use of faba bean caused lower daily
weight gain at the beginning of the growing period and after
adaptation, there were no problems with high proportions of faba
bean in protein concentrate. Then faba bean reached 90% of the
protein concentrate when the control diet included 42% of soybean
meal and 50% of barley. 1.5 kg of that protein concentrate was
allowed per bull and per day. These evaluations established a
general good adaptation of faba bean in ruminant diets and did not
identify any particular quality trait to improve by breeding.
4. Faba beans in human nutrition
Faba beans have been an essential staple food for millennia,
widespread in the Mediterranean area, including continental areas
such as modern Irak, Syria, Iran, and in Northwest India, Pakistan
and Southern China. The high contents in digestible proteins and
starch in their seeds mainly explain this extensive food use; more
recent works have demonstrated the health benefits of faba bean
seeds, because as other pulses they are also good sources of fibres,
and particular vitamins and minerals (Ofuya and Akhidue, 2005;
Champ, 2002). However, their VC constituents may cause health
problems in particular situations.
4.1. Potential toxicity of faba beans
Faba beans are rich in two glucosidic aminopyrimidine
derivatives, V and C, which generate the redox aglycones divicine
(D) (2,6-diamino-4,5-dihydroxypyrimidine) and isouramil (I) (6-
amino-2,4,5-trihydroxypyrimidine), respectively, upon hydrolysis
of the beta-glucosidic bond between glucose and the hydroxyl group
at C-5 on the pyrimidine ring. Faba beans contain high amounts of
ascorbate and varying amounts of L-DOPA glucoside (Arese and De
Flora, 1990). A clear age-dependent distribution of the potentially
hazardous components V, C and ascorbate, was noted in a large
number of faba bean cultivars (Arese and De Flora, 1990). Extremely
high levels of V and C and high levels of ascorbate are present in
young seeds. L-DOPA glucoside was determined in 27 cultivars
showing levels varying from 0.25 to 2.29 g per 100 g of DM (Arese
and De Flora, 1990). L-DOPA is apparently not toxic to G6PD-
deficient subjects (Beutler, 1970; Arese and De Flora, 1990). Faba
bean seeds contain a beta-glucosidase which is able to split the beta-
glucosidic bond of V and C. Its activity is very low in young seeds,
reaches a maximum in ripe seeds and drops again in older seeds
(Arese, 1982; Arese and De Flora, 1990). The faba bean beta-
glucosidase transforms non-toxic V and C into D and I that are toxic
to G6PD-deficient subjects. This enzyme is inactivated by cooking,
seed drying and by acid, such as hydrochloric acid at concentrations
 K. Crépon et al. / Field Crops Research 115 (2010) 329–339
similar to those found in the normal, adult gastric juice. For this
reason, dried and cooked beans are not toxic. G6PD-deficient (see
below) small children and old people are at risk because their gastric
juice is less acidic and the beta-glucosidase of the beans is not
inactivated. D and I are absorbed rapidly and, in association with
ascorbic acid, superstoichiometrically oxidize glutathione (GSH)
contained in millimolar concentration in the RBC. GSH is a very
important intracellular reductant that keeps in the reduced form a
large number of thiol groups localized in enzymes and other proteins
(Arese and De Flora, 1990). When GSH is oxidized, several important
functions of RBC are altered: RBC become rigid, certain essential
membrane proteins are aggregated (for example band 3), important
enzymes are inactivated. As a consequence, RBC are transformed
into senescent RBC and are recognized as non-self cells by the
macrophages of the immune system and rapidly removed (Arese
et al., 2005). In normal RBC, oxidized GSH is rapidly regenerated by a
metabolic cycle of which G6PD is an essential component. In fact,
experiments on normal volunteers have shown that ingestion of
500 g of commercially available faba beans elicited a 30–40% drop in
the GSH level within 3 h from ingestion. However, the original GSH
level was regenerated quickly and the normal volunteers experi-
enced no adverse effects. For this reason, normal individuals may
ingest large amounts of fresh faba beans without any harm.
However, in G6PD-deficient RBC regeneration of oxidized GSH is
extremely slow, D/I become toxic and are thus generally considered
to be the main causative agents of favism. In fact, in G6PD-deficient
subjects D and I liberated from ingested faba beans cause an almost
irreversible oxidation of GSH and, as a consequence, a number of
changes that finally provoke rapid and massive removal of large
numbers of RBC by macrophages (Arese and De Flora, 1990).
4.2. Favism in G6PD-deficient human subjects
Favism is an acute hemolysis caused by the ingestion of faba
beans occurring only in G6PD-deficient human individuals (Arese,
1982; Arese and De Flora, 1990). Hemolysis means that RBC are
destroyed within 24–36 h after faba bean ingestion. A small
portion of destroyed RBC breaks down (lysis) in the blood, but by
far the largest proportion of disappearing RBC is removed by
macrophages. Favism may cause rapid destruction of up to 80% of
circulating RBC and is potentially fatal. Usually favism is benign
and does not require blood transfusion. There is no cure for favism,
except prompt RBC transfusion. Similar hemolytic crisis is also
caused by oxidants (oxidant drugs, such as primaquine, sulfona-
mides; oxidant chemicals ingested by mistake). Vitamin C is
dangerous only if taken in large quantities (>10–20 g). There are
many G6PD variants with low activity. The carriers of the low-
activity Mediterranean and the Chinese Canton variant are more at
risk of favism. The African variant A (20% of normal activity) is not
associated with favism. The association of faba bean consumption
and G6PD deficiency is frequent in the Mediterranean area, the
Middle East (Turkey, Iran and Iraq) and the Far East (Taiwan and
southern China). In Italy, favism is restricted to Sardinia, Sicily and
other Southern Italian regions. Severe favism mostly affects male
children between 2 and 5 years of age (67–75% of the cases of
severe favism), as G6PD deficiency is X-linked and homozygous
females are rare. On average, less than 20% of G6PD-deficient
individuals experience favism in their life, in spite of frequent faba
bean consumption. Favism is usually due to raw fresh seeds (>96%
of all cases), or to fresh seeds incompletely cooked (brief boiling or
frying). Frozen fresh faba beans are as dangerous as fresh beans.
The strong variability in the levels of the various components and
in the beta-glucosidase activity is likely to play a role in triggering
the hemolytic crisis. Small children preferentially eat raw very
young beans (dry weight 15–25%) because they are tender and
sweet. These beans are very high in VC but low in beta-glucosidase.
337
In the recent faba bean lines which carry the gene vc gene, the
content of VC is reduced 10–20-fold in dry mature seeds and 20–30
in fresh seeds (Duc et al., 1989).
4.3. Faba beans, malaria and G6PD deficiency
Today, severe malaria, caused by Plasmodium falciparum is one
of the greatest human afflictions, with 1.5–2.7 deaths and 300–500
millions new cases of clinical illness per year worldwide. It is
estimated that the epidemic spread of falciparum malaria is related
to the development of agriculture within the past 10,000 years
(Spielman et al., 2001; Tishkoff et al., 2001). In this relatively short
period of time, a number of RBC mutations spread in malarious
areas all over the world. Those mutations provide efficient defense
against malaria, and reduce mortality. In the 1930s Haldane and
others introduced the concept of ‘‘balanced polymorphism’’ and
‘‘selective advantage of the heterozygote’’, showing that infectious
diseases like malaria expand the frequency of genes that provide
resistance (Arese, 2006; Tishkoff et al., 2001). Expansion of certain
genes is due either to increase in reproductive rate or decrease in
mortality, ensuing in differential survival of some genotypes at the
loss of others. Heterozygote advantage and balanced polymorph-
ism mean that a mutant allele provides a selective advantage in the
heterozygote but has lethal or severe outcome in the homozygote
(Arese, 2006). In the case of malaria, mutations affecting the RBC
cause disadvantage in the homozygote that is balanced by
protection from malaria in the heterozygote. Thus the frequency
of heterozygotes increases until it is balanced by the negative
effects of the homozygous state (Flint et al., 1993; Haldane,
1949a,b).
Deficiency of G6PD is very widespread. Population studies
indicate 400 million of deficient hemi-heterozygous subjects
worldwide, a likely underestimate (Greene, 1993). Geographic
coincidence between distribution of malaria, G6PD deficiency and
faba beans consumption has been shown in several regions, such as
some Italian regions (Sardinia, Sicily and Calabria), some Medi-
terranean and Middle East countries (Greece, North Africa, Irak,
Iran, Lebanon and Israel), parts of northern India and Pakistan, and
Southern China (Greene, 1993). The general consensus is now in
favor of a protective effect of G6PD deficiency in both males
hemizygotes (about 58% protection) and female heterozygotes
(about 46% protection) (Greene, 1993). Interestingly, it has been
noted that faba bean cultivation and consumption were constantly
coexistent with falciparum malaria and G6PD deficiency. One
possible explanation of this triple coincidence is that consumption
of fresh faba beans, which is maximal during the short rainy season
and coincident with outbursts of seasonal malaria, for example in
Mediterranean countries and in the Middle East, may provide an
additional mechanism of protection against malaria. In fact, D and I
are oxidant species that sensitize early parasite forms, the so-
called ring-forms growing in G6PD-deficient RBC, to be modified
similarly as it occurs in normal RBC senescence and to be removed
by the host immune system at an accelerated rate (Arese, 2006).
5. Conclusions
Most faba bean cultivars produced in the EU have high-tannin
content and their use is limited due to the effect of tannins which
decrease the energy content slightly. Despite this anti-nutritional
factor, T+ faba bean is useful in pig diets where it can be introduced
at up to 350 g/kg. The high protein content of faba bean seeds
makes these seed well adapted to poultry nutrition. When faba
bean is included at levels as high as 250 g/kg in broiler diets, it can
replace soybean meal in a large part. However in different animal
species it has been observed that extreme diets containing
approximately 50% faba beans significantly reduced absorption
338 K. Crépon et al. / Field Crops Research 115 (2010) 329–339
of Zn and Mn (Rubio et al., 1994) and a diet with faba beans as the
only source of protein brought about an impairment in growth,
muscle mass and liver weight in growing rats (Frühbeck et al.,
1999).
Recent genetic improvements have enabled tannins and/or VC
to be highly reduced in faba bean seeds. Removing the tannins
from the seeds increases the energy value and the protein
digestibility of faba bean for pigs and poultry. Removing VC
increases the energy value of faba bean in broiler chicken, and
increases the possible use of faba bean in laying hen diets. For faba
bean, the more promising progress is to remove both tannins and
VC, since the effect of the two anti-nutritional factors on the
nutritional value for broiler chicken is additive.
As far as human consumption is concerned, faba beans are
considered as beneficial. Beside protein and energy supply, effects
of faba beans-enriched diet have been described in humans where
significant decrease in plasma LDL-cholesterol levels were noted
(Frühbeck et al., 1997). Other health benefits are recognized from
their contribution in fibres, particular vitamins and minerals. In
contrast, G6PD-deficient individuals, particularly deficient small
children and old subjects may experience severe RBC destruction
even after consumption of small amounts of fresh beans (Arese and
De Flora, 1990). G6PD deficiency is widespread in large numbers
worldwide due to its protective effect against severe malaria
(Greene, 1993).
The G6PD-deficient individuals once localized in malarial areas
have migrated in large numbers to a number of affluent European,
American or Far-East countries. Those large numbers of G6PD-
deficient subjects are mostly unaware of their genetic defect and
of the potential risk of faba beans consumption, and lack
awareness of the potential danger posed by consumption of
raw faba beans. From this point of view, the availability of faba
beans cultivars with very low levels of VC is a welcome
development brought about by faba bean breeders targeting
the fresh or frozen seed market that should be encouraged by
health authorities.
In addition to the positive role that faba bean seeds can play in
animal and human nutrition that we have reviewed, we underline
that this agronomic species has the potential of several positive
ecological services in production and animal husbandry, which are
described by Köpke and Nemecek (2010).
Acknowledgements
Work funded by the European Union EUFABA project ‘Faba bean
breeding for sustainable agriculture’, QLRT-2001-02307 (FP5-SCP),
by Compagnia di San Paolo, Torino, Italy, by Regione Piemonte,
Ricerca Sanitaria Finalizzata, and by Intramural Funds FIN60 from
University of Torino, Torino, Italy. We thank Dr. R. Thompson for
the English language corrections in this paper.
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