Legume Research

  • Chief EditorJ. S. Sandhu

  • Print ISSN 0250-5371

  • Online ISSN 0976-0571

  • NAAS Rating 6.80

  • SJR 0.391

  • Impact Factor 0.8 (2024)

Frequency :
Monthly (January, February, March, April, May, June, July, August, September, October, November and December)
Indexing Services :
BIOSIS Preview, ISI Citation Index, Biological Abstracts, Elsevier (Scopus and Embase), AGRICOLA, Google Scholar, CrossRef, CAB Abstracting Journals, Chemical Abstracts, Indian Science Abstracts, EBSCO Indexing Services, Index Copernicus
Legume Research, volume 47 issue 5 (may 2024) : 684-694

Advancement in Genomics and Molecular Marker Technologies for Breeding of Faba Bean with Low Vicine-convicine Content: A Review

Sadhan Debnath1,*, Ng. Tombisana Meetei1, Mayank Rai1
1School of Crop Improvement, College of Post-Graduate Studies in Agricultural Sciences, Central Agricultural University, Umiam-793 103, Meghalaya, India.
  • Submitted13-09-2022|

  • Accepted13-12-2022|

  • First Online 27-12-2022|

  • doi 10.18805/LR-5041

Cite article:- Debnath Sadhan, Meetei Tombisana Ng., Rai Mayank (2024). Advancement in Genomics and Molecular Marker Technologies for Breeding of Faba Bean with Low Vicine-convicine Content: A Review . Legume Research. 47(5): 684-694. doi: 10.18805/LR-5041.
Faba bean (Vicia faba L.), is a popular legume crop due to its high protein content (22-38.2%), dietary fibre (12%), medicinal and nutritional values and environmental benefits. It is considered as an excellent source of protein, particularly in developing countries where people cannot afford to buy meat. It also has antioxidant, anti-inflammatory, antiviral, anticancer, anti-diabetic and anti-atherosclerotic effects. But the presence of some key anti-nutritional factors, such as vicine and convicine, tannins, phytic acid etc.  restricts the potential utilization of this crop as food and feed. Vicine and convicine are the thermostable, glucosidic aminopyrimidine derivatives that accumulate in the cotyledons of faba bean during seed development and maturity and cause “Favism” or haemolytic anaemia upon consumption by individuals having deficiency of glucose 6-phosphate dehydrogenase enzyme. As globally millions of people are genetically predisposed to such condition, it is highly desirable to eliminate these compounds from faba bean without compromising yield and other quality characteristics. In this review, we have highlighted the recent advancement in the field of genomics and molecular marker technologies for an easy and efficient selection of faba bean with low vicine and convicine content and the strategies to deploy these efficient tools in the future molecular breeding programs in faba bean.

Faba bean (also broad bean, horse bean or field bean) is an annual herbaceous species of Fabaceae family and is believed to have originated in the Near East (Cubero, 1974). It is considered as a partially crosspollinated species and the rate of outcrossing ranges from 4 to 84% (Bond and Poulsen, 1983). It grows well at temperature about 22°C and tolerates chilling between 0 to 10°C. It is generally cultivated in warm temperate and subtropical countries in the winter and in northern latitudes in the spring (Duc, 1997). Faba bean is widely grown for food and feed as a generous source of high-quality protein, dietary fibre and other valuable nutrients (Duc, 1997). According to the Food and Agriculture Organization Corporate Statistical Database (2019), faba bean is the fourth most widely grown cool season grain legume globally after pea, chickpea and lentil.
Despite of having high yield potential and several nutritional, medicinal and environmental benefits, faba bean cultivation is mostly sporadic and mainly limited to some of the Middle-Eastern and European countries, which is chiefly because of the presence of some anti-nutritional factors (ANFs) and their effects on human and animals’ health. The most potent ANFs limiting its widespread consumption are vicine-convicine (VC) (Duc et al., 1999; Khamassi et al., 2013). Several genomics and molecular studies have been conducted to understand the genetics of the locus responsible for VC and elucidate biosynthetic pathways of these compounds. Further, high-throughput molecular markers flanking the allele for VC have been identified (Tacke et al., 2021; Björnsdotter et al., (2021); Khazaei et al., 2015). This review is focused on understanding the origin and domestication, major challenges in improving production and productivity, nutritional and medicinal values and key ANFs in faba bean. Moreover, it also highlights the achievements in genomics and molecular marker technology for improving production of desired quality faba bean, genetics of VC locus and genes for biosynthesis of VC and recent advances in molecular breeding of faba bean for low vicine-convicine (LVC) content.
Origin and domestication of faba bean
Faba bean is one of the first domesticated food legumes and has a long history of cultivation, started in the early Neolithic times, nearly 8.000 B.C. (Karkanis et al., 2018; Torres et al., 2006; Cubero, 1974). However, the origin and domestication of faba bean is still debated as no wild progenitor of the crop has been discovered yet, or the progenitor may have been extinct (Cubero, 1974; Maxted, 1993). Based on the seed size, faba bean has four subspecies: V. faba major (having large seeds and mainly grown in China and South Mediterranean countries), V. faba equina (having medium seeds and mainly grown in North Africa and Middle Eastern countries), V. faba minor (having small seeds and generally grown in Ethiopia and North Europe) and V. faba paucijuga (which is the primitive form and mainly found from Afghanistan to India) (Cubero, 1974). No successful inter-specific cross between faba bean and other Vicia species has been reported (Duc, 1997). Production of faba bean is mainly concentrated in nine major agro-ecological regions: The Nile valley, Mediterranean basin, Central Asia, East Asia, Ethiopia, Oceania, Latin America, North America and Northern Europe (Torres et al., 2006). It was introduced into India nearly 3000 B.C. through the Mesopotamia probably after the advent of the Arabian spice trade route (Gol, 2015). Since then, faba bean has become a traditional legume crop in the state of Bihar, India. Moreover, this crop is also grown in small scale in Jharkhand, Eastern Uttar Pradesh, Chhatisgarh, Odisha, Madhya Pradesh and some of the North-Eastern states of India (Gol, 2015).
Major challenges in improving production and productivity of faba bean
Global production of faba bean is estimated to be 4.84 million tonnes from an area of 2.5 million ha (FAO Stat, 2019). China is the largest producer of faba bean (37.3%) followed by Ethiopia, Australia, United Kingdom, Germany, France and Egypt. Productivity is the highest in Europe (3.0 tonnes/ha), followed by Asia, Africa, Australia and the Americas (FAO Stat, 2019). Though production of faba bean has increased since last two decades, which may be due to selection and development of high yielding cultivars through various breeding programs, still there is immense need to increase the production and productivity of good quality faba bean varieties to sustain the global food security and nutrition requirement for an ever-increasing population, which is estimated to reach 9.6 billion by 2050 (FAO, 2020). Major faba bean breeding challenges are mainly due to its mixed breeding system, unknown wild progenitor and large genome size of nearly 13 Gb, which is the largest among diploid field crops (Khazaei et al., 2021). Like many other major legume, faba bean yield remains unstable due to biotic and abiotic stresses (Cernay et al., 2015). Moreover, studies reported that, the total grain yield of faba bean is positively correlated with seed protein content, which in turn is determined by genotypes (El- Sherbeeny and Robertson, 1992). However, recent studies have reported that several promising genotypes were identified for seed yield and its component quantitative traits that could be used in different faba bean hybridization programs for yield improvement (Dewangan et al., 2022; Kubure, 2016).

Commercial importance of faba bean
Faba bean could be eaten in several forms such as vegetable fresh, dry seeds, frozen or canned, snacks, stewed broad bean (Medamis), broad bean cakes (Taamia), stewed broad bean paste (Bissara), germinated broad bean soap, thick gruels, purees etc. (Dhull et al., 2021, Pasqualone et al., 2020). In China, Ethiopia, the Middle-East and the Mediterranean, faba bean is used as a breakfast food as soup, stews and paste, whereas, in India, fresh green podsare mainly cooked and consumed as vegetable. Apart from these, faba bean is also widely used as livestock feed for poultry, pigs and horses in many industrialized countries (href="#crépon_2010">Crépon et al., 2010, Guillaume and Bellec, 1977).
Nutritional property
Being an excellent source of protein (22-38.2%), faba bean is mainly used as a cheap source of protein in many developing countries where people find difficult to buy meat (Alghamdi et al., 2012). Its protein content is higher than other common food legumes (Burstin et al., 2011; Griffiths and Lawes, 1978). It is also a good source of carbohydrate (57.3%) with an average starch content of 47%, fibre (12%), lipids (1.2-4.0%), important vitamins (B complex vitamins), bioactive compounds and energy (320 kcal/100 g) (Karaköy et al.,  2018; Baginsky et al., 2013; Ofuya and Akhidue, 2005). Faba bean contains high amount of folic acid (Vitamin B9, 148 mg/100 g), which plays a critical role in synthesis and repair of nucleic acids, amino acid metabolism and prevention of anaemia by helping in production of Red Blood Corpuscles (RBCs) (Singh, 2018). It is also a rich source of mineral elements as it contains good amount of macronutrients like nitrogen (6.40%), phosphorous (0.56%), potassium (1.51%), calcium (0.62%) and magnesium (0.35%) and micronutrients like copper (17.6 mg), zinc (42.7 mg), iron (83.8 mg) and manganese (24.0 mg) per kg (Karaköy et al.,  2018).
Medicinal significance
Flavonoids, tannins, lignins, gallic acid, stillbenes etc. are some of the phenolic compounds present in faba bean. Among these, flavonoids are the most important compounds as they have antioxidant, antiviral, anticancer, anti-inflammatory and anti-atherosclerotic effects (Nijveldt et al., 2001). Faba bean contains L-DOPA (L- 3,4-dihydroxy phenylalanine), which is the precursor of Dopamine (Happiness Hormone) that has the ability to cross blood brain barrier and hence used for treatment of Parkinson’s disease (PD), the second most common neurodegenerative disease in elderly people, leading to disability due to an imbalance between dopamine and acetylcholine in the brain (Topal and Bozoglu, 2016; Oviedo-Silva et al., 2018). The concentration of L-DOPA in dry seeds is nearly 0.07% (Ramya and Thaakur, 2007). It is also a suitable food for diabetic patients, heart and cardiovascular diseases because of its chemical composition (Baginsky et al., 2013).
Genetic constitution and genomic resources
Faba bean is a partially allogamous diploid species with six pairs of remarkably large chromosomes (2 n = 12) and has largest known genome (13 Gbp) among legumes (Sato et al., 2010; Ellwood et al., 2008) and any diploid field crops (Soltis et al., 2003) with more than 85-95 % repetitive DNA (Novák et al.,  2020). The genome of faba bean is about 26, 15.9, 4.0, 3.0, 2.9 times larger than the model legume M. truncatula, Chickpea, Human, Lentil, Pea genomes respectively (Khazaei et al., 2021). Hence, the large genome size highly complicates the identification and location of important agronomic genes as well as the development of saturated linkage maps to be used as tools for Marker Assisted Selection (MAS). Therefore, genomic resources are relatively less advanced in faba bean compared with other grain legume species (Khazaei et al., 2021).
Though initially several genetic maps were developed with the help of morphological characteristics, isozymes, seed protein genes and random amplified polymorphic DNA (RAPD) markers, later faba bean genetic studies and breeding have been enriched due to development of expressed sequence tags (ESTs), microsatellites or simple sequence repeats (SSRs), EST-SSRs, single nucleotide polymorphisms (SNPs) and Kompetitive Allele Specific PCR (KASP) markers (Maalouf et al., 2022; Khazaei et al., 2021; Zanotto et al., 2020; Abuzayed, 2019; Khazaei et al., 2017; Kaur et al., 2014a;). Several mapping populations were developed for flowering, yield-related traits and plant architecture (Avila et al., 2017; Cruz-Izquierdo et al., 2012), biochemical and morphological traits (Ramsay et al., 1995), seed weight (Vaz Patto et al., 1999), drought adaptation-related, morphological traits and vicine-convicine (Khazaei et al., 2015, 2014a, 2014b,), rust resistance (Ijaz, 2018), rust, broomrape and ascochyta blight resistance (Románet_al2004). In the absence of a reference genome assembly for this species, high-throughput approaches such as transcriptome analysis are considered as efficient tools for enrichment of genomic resources.
Antinutritional factors restricting faba bean usage
Key anti-nutritional compounds viz., vicine and convicine, tannin, phytic acid, saponins, lectins (favin) etc. in faba bean seeds can prevent its potential use as a protein source. VC are the thermostable, glucosidic aminopyrimidine derivatives that accumulate in the cotyledons of faba bean during seed development and maturity (Khamassi et al., 2013). The amount of VC in seeds ranges from 3 to 14 g/kg or approx. 0.3 to 1.5% in wild type, however, VC-free genotypes contain only 5-10% (0.3 to 1.4 g/kg) of this amount (Duc et al., 1999). Though VC are present in all parts of faba bean plant, seeds contain these compounds in an approximate 2:1 ratio (Goyoaga et al., 2008).VC are very much unique to the genus Vicia, whereas, Momordica charantia (bitter gourd or bitter melon or bitter apple) is the only species outside this genus containing vicine (Khazaei et al., 2019; Gauttam and Kalia, 2013).
Faba beans with high levels of VC when consumed by human being, may cause a condition called Favism in individuals having deficiency in G6PDactivity. When these compounds are hydrolyzed by β-glucosidase enzyme, they produce aglycones divicine and isouramil, which cause oxidation of glutathione in RBC, resulting in haemolysis of RBC Björnsdotter et al.,  (2021). More than 400 million people (~4% of the world population), upon ingestion of faba bean containing high level of VC, will suffer from favism, which is caused by the human X-chromosomal inherited genetic deficiency of G6PD. Tannins interfere with digestive enzymes by forming complexes with nutrient molecules which results in reduced digestibility (Gutierrez et al., 2007), whereas, phytic acid reduces the bioavailability of minerals (Deshpande and Cheryan, 1984). For improving quality traits in faba bean, major progress on reduction of VC and seed coat tannins, the main ANFs limiting faba bean seed usage, have been recently achieved through gene discovery (Björnsdotteret_al2021; Zanotto et al., 2020; Gutiérrezet_al2020; Gutiérrez and Torres, 2019).
Genetics of vc locus and genes for biosynthesis of VC
Consumption of faba bean is limited by the presence of potent ANF, VC in both fresh and dry seeds. This trait is governed by a locus, designated as VC locus, which has two alleles viz., VC+ (wild type) and vc- (mutant allele). Duc et al., (1989) reported a natural mutant genotype (genebank accession 1268(4) (1), which is a Greek landrace with LVC, after an extensive phenotyping of more than 900 faba bean genotypes. A single recessive allele, vc- confers the LVC phenotype in faba bean. The allele reduces the concentration of VC by more than 95% (10-20-fold reduction) in seeds of faba bean. Homozygosity at VC locus with mutant allele (vc-/vc-) has the potential to alleviate the genetic disorder of favism in G6PD-deficient individuals and prevent dietary disadvantages when such faba beans are used as feed for animals, whereas, heterozygosity leads to intermediate concentration of VC (Tacke et al., 2021; Gallo et al., 2018; Créponet_al2010). Initially, the vc- locus was mapped on chromosome 1 within an interval of 3.6 cM and ~5-10 cM away from the gene for colorless hilum (hc-) (Fig 1A), though this marker doesn’t give much guarantee of LVC without adequate phenotyping (Khazaei et al., 2015; Gutierrez et al., 2006). href="#björnsdotter_2021">Björnsdotteret_al(2021) greatly refined the genetic interval carrying vc- locus to 0.21 cM by employing a population of 1,157 pseudo-F2 individuals from the cross between Hedin/2 (normal phenotype) and Disco/1 (mutant phenotype with low vicine and convicine) (Fig 1B). Very recently, the same locus was fine mapped and the genetic interval was placed to 0.13 cM by employing another set of 58 newly developed SNPs (Tacke et al., 2021) (Fig 1C).

Fig 1: Fragments of chr. 1 of faba bean showing the putative regions of VC locus.

Earlier Duc et al., (1989) emphasized that formation of VC occurs in the seed coat of faba bean and thereafter, these compounds are transported to embryo. Several studies reported that the VC content of faba bean seeds is maternally determined (Khazaei et al., 2017; Khamassi et al., 2013; Ramsay and Griffiths, 1996). Based on a recent gene-to-metabolite correlation study by Björnsdotter et al.,  (2021), it was demonstrated that the contents of these compounds in seed are maternally determined and they are synthesized in maternal tissues and finally transported to embryo. The biosynthetic pathway of these compounds was recently elucidated by Björnsdotter et al.,  (2021), who provided the experimental evidence that, a bifunctional riboflavin biosynthetic protein, RIBA1 plays the principal role in the biosynthesis of VC (Fig 2). The GTP cyclohydrolase II domain of RIBA1, which catalyzes the first step of the riboflavin biosynthetic pathway, also catalyzes the key step in the biosynthesis of VC.

Fig 2: Schematic diagram of biosynthesis of VC in faba bean.

Expression profiling of 20 genes, which were very tightly correlated with vicine revealed that the gene evg_1250620 has the highest expression level in whole seeds during the early seed-filling stage (Björnsdotter et al.,  2021). This gene encodes an isoform of 3,4-dihydroxy-2-butanone-4-phosphate synthase/GTP cyclohydrolase II, which is a bifunctional riboflavin biosynthesis enzyme (designated as VC1). According to Björnsdotter et al.,  (2021), Gene expression profiling revealed that the expression of VC1 was 7.4 times higher in seed coat than embryo. This enzyme activity is very low in young seeds, but at its maximum in matured seeds and again comes down in older seeds (Björnsdotteret_al2021). The enzyme gets inactivated during seed drying process, cooking or by acids treatment similar to adult gastric juice. Their findings clearly suggest that VC are the side products of riboflavin biosynthesis from the purine GTP. They have hypothesized that the cause of the LVC phenotype is a frame shift insertion in the vc1 gene located at vc  allele and simultaneous changes in the amino acid sequence of RIBA1 protein, whose GTP cyclohydrolase II function is destroyed due to the two base pairs (AT) insertional mutation.
Molecular breeding strategy for faba bean with LVC

Faba bean exhibits huge genetic variability for nutritional contents as well as ANFs (Karaköy et al.,  2018; Baginsky et al., 2013; Burstin et al., 2011; Duc et al., 1999). Faba bean genotypes which have relatively low levels of VC, tannins and phytic acids and high phytase activity are beneficial and highly desirable for their potential use in human and animal nutrition. The discovery of the natural mutant with vc locus had led to an increasing interest among plant scientist in breeding and development of LVC faba bean varieties (Duc et al.,1989). But despite of having monogenic inheritance nature of vc- locus, the initial attempts to develop marker assisted selection strategies had been unsuccessful towards the identification of suitable and reliable molecular markers for the vc- allele, which is chiefly because of lack of sequence information and limited genomic data on such a huge and complex genome of this crop (~ 13 Gb; Cooper et al., 2017, Soltis et al., 2003). Consequently, the genomic tools in faba bean are still underdeveloped (Annicchiarico et al., 2017) and therefore, the high syntenic correspondence of this crop to the already sequenced genomes of Medicago truncatula, Cicer arietinum and other legumes of subfamily Faboideae have great importance for genomic analyses (Kaur et al., 2014b, 2012; Cruz-Izquierdo et al., 2012; Ellwood et al., 2008).
Ellwood et al., (2008) for the first time provided a sequence-based genetic map of Vicia faba, which allowed the global pattern of Vf -Mt synteny to be observed. But the most clear and accurate picture of macrosynteny between Vf and Mt comes from the study of Webb et al., (2016). A single consensus map containing 687 SNPs and six linkage groups was constructed by merging six linkage maps. This sequence-based consensus map has been utilized for exploring synteny with the most closely related legume species such as Medicago truncatula, Cicer arietinum, Lens culinaris etc. (O’Sullivan and Angra, 2016). These SNPs are valuable tools for genotyping unexplored faba bean genotypes and synteny based trait targeting.
As the genetic interval carrying vc- allele is present at the distal end of the short arm of chromosome 1 of faba bean, the tightly linked high-throughput markers flanking the allelic position are of utmost importance for an efficient marker assisted selection (Webb et al., 2016). Although, initially Khazaei et al., (2015) genotyped a segregating population of 210 F5 recombinant inbred lines with a set of 188 SNPs and identified a strong single QTL determining VC concentration on chromosome 1, flanked by markers 1.0 cM upstream and2.6 cM downstream of the QTL (Fig 1A), but unfortunately these markers turned out to be non-diagnostic for LVC in a large set of germplasm. For the first time, Khazaei et al., (2017) developed and validated a reliable, low-cost, breeder-friendly, robust and high-throughput KASP marker system (KASP_vcp2) for large scale screening of LVC faba bean. This marker system is a highly efficient tool, because it can detect individual homozygous or heterozygous plants very easily without any phenotyping for VC. Hence, this marker has the potential to achieve rapid homozygosity at VC locus for LVC (vc-/vc-) (Khazaei et al., 2017) and facilitate any faba bean breeding program through rapid acceleration in the process of genetic improvement in faba bean.  
In another attempt, Björnsdotter et al.,  (2021) narrowed down the interval for vc- allele by employing SNPs from a previous map based on 3.6 cM interval reported by Khazaei et al., (2015) and individual SNPs designed based on markers mined from RNA-seq data. The SNPs (AX-181184219/AX-181160542 and AX-181438475), are flanking the genetic interval carrying VC locus, spanning a genetic distance of 0.21 cM (Fig 1B). The interval also contains the KASP marker (vcp2) previously found to be tightly linked with VC gene as reported by Khazaei et al., (2017). Recently, the same locus was fine mapped to a region of 0.13 cM by using 58 newly developed SNPs (Tacke et al., 2021). Markers VFS18002.A51 and VFS18002.A54, which span an interval of 0.13 cM are reported to be the final boundary markers which define the core region where the VC locus must be located (Fig 1C). They also reported that, based on the data generated by Song, (2017) and Khazaei et al., (2017), markers SNP384, Vf_Mt2g009320_001 and Vf_Mt2g010740_001 and their chromosomal vicinity deemed to be the most likely region which contains the VC gene. These markers could be used as potential diagnostic marker for VC content in faba bean.

Faba bean, a highly proteinaceous underutilized legume, has immense potential to fulfil the nutritional requirement and maintain general wellbeing and health of human beings including large number of low-income families of several states of India. High yielding faba bean varieties with improved tolerance to several biotic and abiotic stresses were released in different parts of the world. However, food safety concerns remain as the potential impediment, which is threatening human and animal health and delaying large scale cultivation and commercialization of this crop as food and feed for nutrition. Recent discovery of the VC1 gene and fine mapping of VC locus using tightly linked SNPs will greatly facilitate the selection of faba bean for LVC. Tightly linked SNP markers mentioned here could serve as highly useful tools to explore LVC content in large number of local and unadopted germplasms of major faba bean growing regions including North-East India. The significant SNPs near close vicinity to the VC locus or residing at the VC locus could be useful for high throughput SNP genotyping for low content of VC using KASP-assay. Reliable SNPs, which show adequate association with VC phenotyping can also be characterized by using ARMS markers for tetra-primer Amplification Refractory Mutation System PCR (ARMS-PCR) assay. One-step ARMS-PCR is highly efficient and cost effective as this assay could easily identify the genotype of a plant based on homozygous or heterozygous status of an SNP allele. Moreover, a PCR reaction volume of 10µl is generally sufficient and alleles are easily separated on simple agarose gel. SSRs or microsatellites markers, which were found to be linked with VC content in faba bean, could serve as valuable tools for screening faba bean genotypes for low and high VC content in a limited resourced facility. However, further study needs to be done with these microsatellites markers for generating conclusive data in faba bean. As the sequence of the VC locus is publicly available now, breeders can easily design primers specific to the two bp insertional mutation, which has led to a substantial reduction in VC content in faba bean. Presently, several collaborative reference genome assembly and pan-genome initiatives are underway and these are expected to provide annotated reference transcriptomes for faba bean and hence, accelerate high-throughput genotyping in this crop. Advanced faba bean genotypes with low vicine and convicine could efficiently be used in future breeding programmes in faba bean worldwide to prevent the metabolic food disorder of Favism in susceptible individuals.
 All authors contributed equally in literature search, compiling writing and editing.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

  1. Abuzayed, M.A. (2019). Molecular genetic analysis in faba bean (Vicia faba L.). Ph.D. thesis submitted to the Graduate School of Engineering and Sciences of İzmir Institute of Technology.

  2. Alghamdi, S.S., Migdadi, H.M., Ammar, M.H., Paull, J.G. and Siddique, K.H.M. (2012). Faba bean genomics: Current status and future prospects. Euphytica. 186: 609-624.

  3. Annicchiarico, P., Nazzicari, N., Wei, Y., Pecetti, L. and Brummer, E.C. (2017). Genotyping by sequencing and its exploitation for forage and cool-season grain legume breeding. Frontiers in Plant Science. 8: 679. DOI: 10.3389/ fpls.2017.00679.

  4. Avila, C.M., Ruiz-Rodríguez, M.D., Cruz-Izquierdo, S., Atienza, S.G., Cubero, J.I. and Torres, A.M. (2017). Identification of plant architecture and yield-related QTL in Vicia faba L. Molecular Breeding. 37(7): 88. DOI:10.1007/s11032-017-0688-7.

  5. Baginsky, C., Pen˜a-Neira, A., Ca´ceres, A., Herna´ndez, T., Estrella, I., Morales, H. and Pertuze, R. (2013). Phenolic compound  composition in immature seeds of faba bean (Vicia faba L.) varieties cultivated in Chile. Journal of Food Composition  and Analysis. 31(1): 1-6.

  6. Björnsdotter, E., Nadzieja, M., Chang, W., Escobar-Herrera, L., Mancinotti, D., Angra, D., Xia, X. et al. (2021). VC1 catalyzes  a key step in the biosynthesis of vicine in faba bean. Nature Plants. 7: 923-931.

  7. Bond, D.A. and Poulsen, M.H. (1983). Pollination. In: [Hebblethwaite, P.D. (Ed.)], The Faba Bean (Vicia faba L.). Butterworths, London, UK. 77-101.

  8. Burstin, J., Gallardo, K., Mir, R.R., Varshney, R.K. and Duc, G. (2011). Improving Protein Content and Nutrition Quality (Chapter 20). In: Biology and Breeding of Food Legumes. [Pratap, A. and Kumar, J. (Eds.)], New Delhi, India: CABI. 314-328. https://doi.org/10.1079/ 9781845937669.0000.

  9. Cernay, C., Ben-Ari, T., Pelzer, E., Meynard, J.M. and Makowski, D. (2015). Estimating variability in grain legume yields across Europe and the Americas. Scientific Reports. 5: 11171. https://doi. org/10.1038/sre p11171.

  10. Cooper, J.W., Wilson, M.H., Derks, M.F.L., Smit, S., Kunert, K.J., Cullis, C. and Foyer, C.H. (2017). Enhancing faba bean (Vicia faba L.) genome resources. Journal of Experimental  Botany. 68: 1941-1953. https://doi.org/10.1093/jxb/erx117.

  11. Crépon, K., Marget, P., Peyronnet, C., Carrouée, B., Arese, P. and Duc, G. (2010). Nutritional value of faba bean (Vicia faba L.) seeds for feed and food. Field Crop Research. 115: 329-339.

  12. Cruz-Izquierdo, S., Avila, C.M., Satovic, Z., Palomino, C., Gutierrez, N., Ellwood, S.R., Phan, H.T.T., Cubero, J.I. and Torres, A.M. (2012). Comparative genomics to bridge Vicia faba with model and closely-related legume species: Stability of QTLs for flowering and yield-related traits. Theoretical and Applied Genetics. 125: 1767-1782.

  13. Cubero, J.I. (1974). On the evolution of Vicia faba L. Theoretical and Applied Genetics. 45(2): 47-51.

  14. Deshpande, S. and Cheryan, M. (1984). Effect of phytic acid, divalent  cations and their interactions on á-amylase activity. Journal of Food Science. 49: 516-519.

  15. Dewangan, N.K., Dahiya, G.S., Janghel, D.K. and Dohare, S. (2022). Diversity analysis for seed yield and its component traits among faba bean (Vicia faba L.) germplasm lines. Legume  Research. 45(6): 689-694. DOI: 10.18805/LR-4301.

  16. Dhull, S.B., Kidwai, M.K., Noor, R., Chawla, P. and Rose, P.K. (2021). A review of nutritional profile and processing of faba bean (Vicia faba L.). Legume Science. e129. https://doi.org/ 10.1002/leg3. 129.

  17. Duc, G. (1997). Faba bean (Vicia faba L.). Field Crops Research. 53: 99-10.

  18. Duc, G., Marget, P., Esnault, R., Guen, J.L. and Bastianelli, D. (1999). Genetic variability for feeding value of faba bean seeds (Vicia faba) comparative chemical composition of isogenics involving zero-tannin and zero-vicine genes. Journal of Agricultural Science. 133(2): 185-196.

  19. Duc, G., Sixdenier, G., Lila, M. and Furstoss, V. (1989). Search of Genetic Variability for Vicine and Convicine Content in Vicia faba L.: A First Report of a Gene Which Codes for  Nearly Zero vicine and Zero convicine Contents. In: Recent Advances of Research in Antinutritional Factors in Legume Seeds; [Huisman, J., van der Poel, A.F.B., Liener, I.E., (Eds.)]; Wageningen Academic Publishers: Wageningen, The Netherlands. pp 305-313.

  20. Ellwood, S.R., Phan, H.T.T., Jordan, M., Hane, J., Torres, A.M., Avila, C.M., Cruz-Izquierdo, S. and Oliver R.P. (2008). Construction of a comparative genetic map in faba bean (Vicia faba L.); conservation of genome structure with Lens culinaris. BMC Genomics. 9: 1-11. doi: 10.1186/1471-2164-9-380.

  21. El-Sherbeeny, M.H. and Robertson, L.D. (1992). Protein content variation in a pure line faba bean (Vicia faba) collection. Journal of the Science of Food and Agriculture. 58(2): 193-196. https://doi. org/10.1002/ (ISSN) 1097-0010.

  22. Food and Agriculture Organization Corporate Statistical Database. (2019). Food and Agriculture Organization of the United Nations, Available Online: faostat.fao.org. 

  23. Food and Agriculture Organization. (2020). The state of food security and nutrition in the world. Transforming Food Systems for Affordable Healthy Diets.

  24. Gallo, V., Skorokhod, O.A., Simula, L.F., Marrocco, T., Tambini, E., Schwarzer, Marget, P., Duc, G. and Arese, P. (2018). No red blood cell damage and no hemolysis in G6PD-deficient  subjects after ingestion of low vicine/convicine Vicia faba seeds. Blood. 131(14): 1621-1625. DOI: 10.1182/blood- 2017-09-806364.

  25. Gauttam, V., Kalia, A.N. (2013). High performance thin layer chromatography method for simultaneous estimation of vicine, trigonelline and withaferin-a in a polyherbal antidiabetic  formulation. International Journal of Pharmacy and Pharmaceutical Sciences. 5(2): 367-371.

  26. Gol, S. (2015). Determination of genetic diversity and population structure in faba bean (Vicia faba L.). Thesis Submitted to the Graduate School of Engineering and Sciences of İzmir Institute of Technology. http://hdl.handle.net/11147/4342.

  27. Goyoaga, C., Burbano, C., Cuadrado, C., Varela, A., Guillamón, E., Pedrosa, M.M., Muzquiz, M. (2008). Content and distribution of vicine, convicine and l-DOPA during germination and seedling growth of two Vicia faba L. varieties. European Food Research and Technology. 227(5): 1537-1542. DOI: 10.1007/s00217-008-0876-0.

  28. Griffiths, D.W. and Lawes, D.A. (1978). Variation in the crude protein content of field beans (Vicia faba L.) in relation to the possible improvement of the protein content of the crop. Euphytica. 27(2): 487-495. https://doi.org/10.1007/ BF00043174.

  29. Guillaume, J. and Bellec, R. (1977). Use of field beans (Vicia faba L.) in diets for laying hens. British Poultry Science. 18(5): 573-583. DOI: 10.1080/00071667708416406.

  30. Gutiérrez, N. and Torres, A.M. (2019). Characterization and diagnostic marker for TTG1 regulating tannin and anthocyanin biosynthesis in faba bean. Scientific Report. 9: 16174.

  31. Gutiérrez, N., Avila, C.M. and Torres, A.M. (2020). The bHLH transcription factor VfTT8 underlies zt2, the locus determining zero tannin content in faba bean (Vicia faba L.). Scientific Report. 10: 14299.

  32. Gutierrez, N., Avila, C.M., Duc, G., Marget, P., Suso, M.J., Moreno, M.T. and Torres, A.M. (2006). CAPs markers to assist selection for low vicine and convicine contents in faba bean (Vicia faba L.). Theoretical and Applied Genetics. 114: 59-66.

  33. Gutierrez, N., Avila, C.M., Rodriguez-Suarez, C., Moreno, M.T. and Torres, A.M. (2007). Development of SCAR markers linked to a gene controlling absence of tannins in faba bean. Molecular Breeding. 19: 305-314.

  34. Ijaz, U. (2018). Molecular Mapping and Microscopic Analysis of Faba Bean Uromyces Viciae-fabae Host-pathogen Interaction. PhD Thesis, The University of Sydney, Sydney, Australia.

  35. Karaköy, T., Demirbas, A., Toklu F., Gürsoy, N., Tugay Karagöl, E., Uncuer, D. and Ozkan, H. (2018). Assessment of micro and macro nutrients contents in the Turkish faba bean germplasm. Agriculture for Life, Life for Agriculture Conference Proceedings. 1(1): 72-78.

  36. Karkanis, A., Ntatsi, G., Lepse, L., Fernández, J.A., Vågen, I.M., Rewald, B., Alsiòa, I., Kronberga, A., Balliu, A., Olle, M., Bodner, G., Dubova, L., Rosa, E. and Savvas, D. (2018). Faba bean cultivation - revealing novel managing practices for more sustainable and competitive european cropping systems. Frontiers in Plant Science. 9: 1115. doi: 10.3389/fpls.2018.01115. PMID: 30116251; PMCID: PMC6083270.

  37. Kaur, S., Cogan N.O.I., Forster J.W. and Paull J.G. (2014a). Assessment  of genetic diversity in faba bean based on single nucleotide  polymorphism. Diversity. 6(1): 88-101.

  38. Kaur, S., Kimber, R.B.E., Cogan, N.O.I., Materne, M., Forster, J.W. and Paull, J. (2014b). SNP discovery and high-density genetic mapping in faba bean (Vicia faba L.) permits identification of QTLs for ascochyta blight resistance. Plant Science. 217-218: 47-55.

  39. Kaur, S., Pembleton, L., NOI, C., Savin, K, Leonforte, T., Paull, J., Materne, M., Forster, J. (2012). Transcriptome sequencing  of field pea and faba bean for discovery and validation of SSR genetic markers. BMC Genomics. 13: 104. https:// doi.org/10.1186/1471-2164.

  40. Khamassi, K., Jeddi, F.B., Hobbs, D., Irigoyen, J., Stoddard, F., O’Sullivan, D.M. and Jones, H. (2013). A baseline study of vicine-convicine levels in faba bean (Vicia faba L.) germplasm. Plant Genetic Resources: Characterisation and Utilization. 11(3): 250-257.

  41. Khazaei, H., O’Sullivan, D.M., Jones, H., Pitts, N., Sillanpää, M.J., Pärssinen, P., Manninen, O. and Stoddard, F.L. (2015). Flanking SNP markers for vicine-convicine concentration in faba bean (Vicia faba L.). Molecular Breeding. 35(1): 38. https://doi.org/10.1007/s11032-015-0214-8.

  42. Khazaei, H., O’Sullivan, D.M., Sillanpää, M.J. and Stoddard, F.L. (2014a). Use of synteny to identify candidate genes underlying QTL controlling stomatal traits in faba bean (Vicia faba L.). Theoretical and Applied Genetics. 127: 2371-2385.

  43. Khazaei, H., O’Sullivan, D.M., Sillanpää, M.J. and Stoddard, F.L. (2014b). Genetic analysis reveals a novel locus in Vicia faba decoupling pigmentation in the flower from that in the extra-floral nectaries. Molecular Breeding. 34: 1507-1513.

  44. Khazaei, H., O’Sullivan, D.M., Stoddard, F.L., Adhikari, K.N., Paull, J.G., Schulman, A.H. andersen, S.U. and Vandenberg, A. (2021). Recent advances in faba bean genetic and genomic tools for crop improvement. Legume Science. 3(3): e75. https://doi.org/10.1002/leg3.75.

  45. Khazaei, H., Purves, R.W., Song, M., Stonehouse, R., Bett, K.E., Stoddard, F.L. and Vandenberg, A. (2017). Development and validation of a robust, breeder-friendly molecular marker for the vc- locus in faba bean. Molecuar Breeding. 37: 140. 10.1007/s11032-017-0742-5.

  46. Khazaei, H., Subedi, M., Nickerson, M., Martínez-Villaluenga, C., Frias, J. and Vandenberg, A. (2019). Seed protein of lentils: Current status, progress and food applications. Foods (Basel, Switzerland). 8(9). DOI: 10.3390/foods8090391.

  47. Kubure, T.E., Raghavaiah, C.V. and Hamza, I. (2016). Production potential of faba bean (Vicia faba L.) genotypes in relation to plant densities and phosphorus nutrition on Vertisols of Central Highlands of West Showa Zone, Ethiopia, East Africa. Advances in Crop Science and Technology. 4: 214. doi:10.4172/2329-8863.1000214.

  48. Maalouf, F., Abou-Khater, L., Babiker, Z., Jighly, A., Alsamman, A.M., Hu, J., Ma, Y., Rispail, N., Balech, R., Hamweih, A., Baum, M. and Kumar, S. (2022). Genetic dissection of heat stress tolerance in faba bean (Vicia faba L.) using GWAS. Plants. 11(9): 1108. https://doi.org/10.3390/ Plants. 11091108.

  49. Maxted, N. (1993). A phenetic investigation of Vicia L. subgenus Vicia (Leguminosae, Vicieae). Botanical Journal of The Linnean Society. 111: 155-182.

  50. Nijveldt, R.J., van Nood, E., van Hoorn, D.E., Boelens, P.G., Norren, K.V. and van Leeuwen, P.A. (2001). Flavonoids: A review of probable mechanisms of action and potential applications.  American Journal of Clinical Nutrition. 74(4): 418-25.

  51. Novák, P., Guignard, M.S., Neumann, P., Kelly, L.J., Mlinarec, J., Koblížková, A., Dodsworth, S., Kovaøík, A., Pellicer, J., Wang, W., Macas, J., Leitch, I.J. and Leitch, A.R. (2020). Repeat-sequence turnover shifts fundamentally in species  with large genomes. Nat. Plants. 6: 1325-1329.

  52. O’Sullivan, D.M. and Angra, D. (2016). Advances in faba bean genetics and genomics. Frontiers in Genetics. 7: 150. doi: 10.3389/fgene.2016.00150.

  53. Ofuya, Z.M. and Akhidue, V. (2005). The role of pulses in human nutrition: A review. Journal of Applied Sciences and Environmental Management. 9(3): 99-104.

  54. Oviedo-Silva, C.A., Elso-Freudenberg, M. and Aranda-Bustos, M. (2018). L-DOPA trends in different tissues at early stages of Vicia faba growth: Effect of tyrosine treatment. Applied Sciences. 8: 2431. doi:10.3390/app8122431.

  55. Pasqualone, A., Abdallah, A. and Summo, C. (2020). Symbolic meaning and use of broad beans in traditional foods of the Mediterranean Basin and the Middle East. Journal of Ethnic Food. 7: 39. https://doi.org/10.1186/s42779-020- 00073-1.

  56. Ramsay, G. and Griffiths, D.W. (1996). Accumulation of vicine and convicine in Vicia faba and V. Narbonensis. Phytochemistry.  42(1): 63-67. 

  57. Ramsay, G., van de Ven, W., Waugh, R., Griffiths, D.W. and Powel, W. (1995). Mapping Quantitative Trait Loci in Faba Beans. In: Improving Production and Utilization of Grain Legumes. [AEP (Ed.)], Copenhagen, Denmark. 2nd European conference on grain legumes. pp 444–445.

  58. Ramya, K.B. and Thaakur, S. (2007). Herbs containing L- Dopa: An update. Ancient Science of life. 27(1): 50-55.

  59. Román, B., Satovic, Z., Pozarkova, D., Macas, J., Dolezel, J., Cubero, J.I. and Torres, A.M. (2004). Development of a composite map in Vicia faba L. breeding applications and future prospects. Theoretical and Applied Genetics. 108: 1079-1088.

  60. Sato, S., Isobe, S. and Tabata, S. (2010). Structural analyses of the genomes in legumes. Current Opinion in Plant Biology. 13(2): 146-152.

  61. Singh, J. (2018). Folate content in legumes. Biomedical Journal of Scientific and Technical Research. 3(4): 1-6.

  62. Soltis, D.E., Soltis, P.S., Bennett, M.D. and Leitch, I.J. (2003). Evolution of genome size in the angiosperms. Amrican Journal of Botany. 90: 1596-1603.

  63. Song, M. (2017). Preventing favism by selecting faba bean mutants using molecular markers.  STEM Fellowship Journal. 3(1): 2-6. DOI: 10.17975/sfj-2017-001.

  64. Tacke, R., Ecke, W., Höfer, M., Sass, O. and Link, W. (2021). Zooming into the genomic vicinity of the major locus for vicine and convicine in faba bean (Vicia faba L.). Bio Rxiv. 2021.02.19.431996. https://doi.org/10.1101/2021. 02.19.431996.

  65. Topal, N. and Bozoðlu, T. (2016). Determination of L-dopa (l-3, 4- dihydroxyphenylalanine) content of some faba bean (Vicia faba L.) genotypes. Journal of Agricultural Sciences. 22(2): 145-151.

  66. Torres, A.M., Román, B., Avila, C.M., Satovic, Z., Rubiales, D., Sillero, J.C., Cubero, J.I. and Moreno, M.T. (2006). Faba bean breeding for resistance against biotic stresses: Towards application of marker technology. Euphytica. 147(1-2): 67-80. doi:10.1007/s10681-006-4057-6.

  67. Vaz Patto, M.C., Torres, A.M., Koblizkova, A., Macas, J. and Cubero, J.I. (1999). Development of a genetic composite map of Vicia faba using F2 populations derived from trisomic plants. Theoretical and Applied Genetics. 98: 736-743.

  68. Webb, A., Cottage, A., Wood, T., Khamassi, K., Hobbs, D., Gostkiewicz,  K., White, M., Khazaei, H. et al. (2016). A SNP-based consensus genetic map for synteny-based trait targeting in faba bean (Vicia faba L.). Plant Biotechnology Journal. 14: 177-185. doi:10.1111/pbi.12371.

  69. Zanotto, S., Vandenberg, A. and Khazaei, H. (2020). Development and validation of a robust KASP marker for zt2 locus in faba bean (Vicia faba). Plant Breeding. 139: 375-380.

Editorial Board

View all (0)