# Drought and heat affect common bean minerals and human diet-What we know and where to go

Alessia Losa, Juan Vorster, Eleonora Cominelli, Francesca Sparvoli, Dario Paolo, Tea Sala, Marika Ferrari, Marina Carbonaro, Stefania Marconi, Emanuela Camilli, et al.

► To cite this version:

Alessia Losa, Juan Vorster, Eleonora Cominelli, Francesca Sparvoli, Dario Paolo, et al., Drought and heat affect common bean minerals and human diet-What we know and where to go. Food and Energy Security, 2022, 11 (1), pp.e351. 10.1002/fes3.351. hal-03781802

HAL Id: hal-03781802

https://hal.inrae.fr/hal-03781802

Submitted on 20 Sep 2022

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>
# Drought and heat affect common bean minerals and human diet—What we know and where to go

Alessia Losa<fnref n="1" /> | Juan Vorster<fnref n="2" /> ID | Eleonora Cominelli<fnref n="3" /> | Francesca Sparvoli<fnref n="3" /> |

Dario Paolo<fnref n="3" /> | Tea Sala<fnref n="1" /> | Marika Ferrari<fnref n="4" /> | Marina Carbonaro<fnref n="4" /> |

Stefania Marconi<fnref n="4" /> | Emanuela Camilli<fnref n="4" /> | Emmanuelle Reboul<fnref n="5" /> | Boaz Waswa<fnref n="6" /> |

Beatrice Ekesa<fnref n="6" /> | Francisco Aragão<fnref n="7" /> | Karl Kunert<fnref n="2" /> ID

## Correspondence

Karl Kunert, Department Plant and Soil Sciences, Forestry and Agricultural Biotechnology Institute, University of Pretoria, Hillcrest, Pretoria 0002, South Africa.

Email: karl.kunert@up.ac.za

## Funding information

ERA-NET co-funding on Food Systems and Climate (FOSC) BIO-BELIEF project (Reference Number: FOSC-288)

## Abstract

Global climate change, causing large parts of the world to become drier with longer drought periods, severely affects production of common beans (*Phaseolus vulgaris* L.). The bean is worldwide the most produced and consumed food grain legume in the human diet. In common beans, adapted to moderate climates, exposure to drought/heat stress not only results in significant reduction of bean yield but also the nutritional value. This review explores the contribution of common beans to food and nutrient security as well as health. Also discussed is the existing knowledge of the impact of drought/heat stress, associated with a changing climate, specifically on iron (Fe) and phytic acid (PA) that are both among the most important mineral and anti-nutritional compounds found in common beans. Further discussed is how the application of modern “omics” tools contributes in common beans to higher drought/heat tolerance as well as to higher Fe and reduced PA content. Finally, possible future actions are discussed to develop new common bean varieties with both improved drought/heat tolerance and higher mineral (Fe) content.

## KEYWORDS

antinutrients, biofortification, climate change, common bean, drought/heat, legumes, minerals, *Phaseolus vulgaris*, phytic acid, pulses

<figure><img src="image_2.png" /></figure>

<footnote n="1">¹Council for Research in Agriculture and Economics, Research Centre for Genomics and Bioinformatics (CREA-GB), Montanaso, Italy</footnote>

<footnote n="2">²Department Plant and Soil Sciences, Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria, South Africa</footnote>

<footnote n="3">³National Research Council, Institute of Agricultural Biology and Biotechnology (CNR-IBBA), Milan, Italy</footnote>

<footnote n="4">⁴Council for Agricultural Research and Economics, Research Centre for Food and Nutrition (CREA-AN), Rome, Italy</footnote>

<footnote n="5">⁵Aix-Marseille University, INRAE, INSERM, C2VN, Marseille, France</footnote>

<footnote n="6">⁶International Center for Tropical Agriculture (CIAT), CIAT Regional Office for Africa, Nairobi, Kenya</footnote>

<footnote n="7">⁷Embrapa Recursos Genéticos e Biotecnologia, Norte, Brazil</footnote>
# 1 | INTRODUCTION

## 1.1 | Climate change and crop production

Global climate change will severely affect the UN goal to sustainably produce enough food by 2050 to feed a projected global population of 9.1 billion people. This is despite previous progress in addressing global undernutrition with increased food production by agricultural expansion and intensification (Myers et al., 2017). Global climate change, which has a long-term impact in the form of different abiotic stresses (Redden, 2013), will particularly cause large parts of the world to become drier with longer drought periods, more intense heat and irregular rainfall. These changes will severely affect agriculture as well as the stability and distribution of food production (Kellogg & Schware, 2019). These changes in the climatic conditions might even become progressively worse (Hao et al., 2018). A major factor in driving such temperature increase and altering precipitation patterns is a higher atmospheric carbon dioxide (CO₂) concentration (Kellogg & Schware, 2019; Lindsey, 2020).

Warmer and drier climatic conditions, resulting in increased evaporative losses, will also drive the demand for more water. A substantial part of the world's agriculture is, however, already suffering from chronic soil water shortages due to severe drought conditions (Nadeem et al., 2019). Predicted shifts in crop production, due to regional changes in temperature and rainfall patterns, might further worsen local food shortages. The future demand for affordable nutritious food will, therefore, require integration of such regional predictions within drought monitoring and forecasting (Leisner, 2020; Mbiriri et al., 2018; Myers et al., 2017; Parsons et al., 2019; Zolina et al., 2013).

## 1.2 | Drought/heat stress and plant growth

Stresses associated with a changing climate are predicted to severely impact plant metabolism as well as soil fertility and carbon sequestration. This impact will limit plant growth and productivity and, ultimately, availability of nutritious food (Dankher & Foyer, 2018; Myers et al., 2017). A higher atmospheric CO₂ concentration affects, for example, the nutritional composition of crops by reducing the protein content of the edible plant parts and also lowering concentrations of important minerals (Loladze, 2014; Medek et al., 2017; Myers et al., 2014). In particular, soil water deficit, due to drought/heat conditions, causes considerable reduction in plant productivity. This reduction is specifically evident in areas where the agricultural

system is dependent on rainfall, such as on the Southern Plains of the USA and in eastern Africa (Adhikari et al., 2015; Ahmed & Stepp, 2016; Steiner et al., 2018). Impacts of drought/heat range from negatively affecting all plant development stages with key physiological, biochemical and metabolic pathways seriously disrupted (Dankher & Foyer, 2018). Temperature extremes are further more strongly associated with a reduction in crop yield, compared to precipitation extremes, and irrigation partly limits the negative effects of high temperatures (Vogel et al., 2019). While plants are always exposed to a combination of stresses under field conditions (Hussain et al., 2018), most studies have focused on the impact of individual stresses on crop yield (Fahad et al., 2017). Combined drought and high temperature stress is known to reduce yields of maize, soybean and wheat (Matiu et al., 2017). Physiological characterization of plants exposed to either drought or heat stress, or a combination of both stresses, has indicated that combined stresses have several distinctive characteristics. These characteristics include low photosynthesis combined with increased respiration and closed stomata combined with increased leaf temperature (Mittler, 2006). However, how plants respond to a combination of different abiotic stresses cannot be simply inferred by the response of the plant to an individual stress (Mittler, 2006). To develop strategies maintaining crop productivity under such individual or combined stresses is, therefore, a major future research challenge (Ferguson, 2019).

## 1.3 | Potential of legumes

Legumes, as members of the large Fabaceae (Leguminosae) family, are predominantly grown in the world's tropical and subtropical areas. The production and consumption of pulses, the edible seeds of legumes, has further greatly increased over the last 15 years (Rawal & Navarro, 2019). Legumes include a number of important agricultural and food plants, such as Glycine max (soybean) and Phaseolus species (beans). They have an important function in both the diversification and sustainable intensification of agriculture. Apart from being a major dietary protein source, legumes are a rich source of minerals to humans and animals. Stresses, associated with changing climatic conditions, pose, however, a severe threat towards their growth, yield potential as well as nutritional value (Foyer et al., 2016; Gepts et al., 2008; Latef & Ahmad, 2015; Nadeem et al., 2019; Sica et al., 2021; Vasconcelos et al., 2020). An important feature of legumes is further the ability to form root nodules allowing to fix atmospheric nitrogen. With the help of rhizobia, legumes reduce atmospheric nitrogen to ammonia in these root nodules with various genes
required for symbiotic nitrogen fixation (SNF) (Roy et al., 2020). The natural ability to add fixed nitrogen into soils renders legumes a keystone species for natural and agricultural ecosystems, injecting over 50 million tons of nitrogen into the soil per annum (Roy et al., 2020). SNF, which significantly contributes not only to protein production but also to increase in soil fertility (Sørensen & Sessitsch, 2007; Wagner, 2011), is, however, highly drought sensitive (Kunert et al., 2016).

## 1.4 | Common beans and abiotic stress

Among the legumes, common bean (*Phaseolus vulgaris* L.; Figure 1), which is the focus of this review, is the most produced and consumed food grain legume worldwide. Common beans are adapted to relatively moderate climatic conditions and not to extreme climatic and edaphic environments. Day temperature exceeding 30°C or night temperatures higher than 20°C can significantly reduce seed yield due to flower abscission, development of parthenocarpic pods (pin pods), lower seed set per pod as well as decreased seed size (Rainey & Griffiths, 2005). Soil degradation and factors, such as soil acidity and aluminium toxicity, lead to deficiencies in nitrogen and phosphorus. This can limit root development and consequently access to soil moisture (Samago et al., 2018).

Growth of legumes is generally affected by several abiotic stresses causing as a response changes in the plant phenotype (Araújo et al., 2015). Among the reported changes in common beans due to abiotic stresses are less leaf expansion due to ultraviolet-B radiation (Riquelme et al., 2007), increase of production of malondialdehyde, antioxidants and carotenoids, a decrease in the chlorophyll content of leaves due to heavy metal exposure (Zengin, 2013), reduction of seedling emergence after low temperature treatment (De Ron et al., 2016), as well as suppression of growth, photosynthesis and transpiration under high soil salinity (Kaymakanova et al., 2008). Changes in protein expression, reduced germination, stunted growth, serious damage to photosynthesis and

a reduction in nutrient uptake particularly occur after exposure to drought/heat stress, (Mathobo et al., 2017; Nadeem et al., 2019; Zadražnik et al., 2013). In addition, a shallow root system further renders common beans vulnerable to any shortage of soil water with the reproductive stage, which includes flowering and pod-filling, also very drought-sensitive (Daryanto et al., 2017).

Adaptation, particularly to drought conditions, includes the improvement in the photosynthetic capacity, water-use efficiency and adaptation to different environments. Such adaptation has been already found in a few common bean genotypes, including the BAT477 race (Beebe et al., 2008; Polania, Rao, et al., 2016), and selection for drought resistance improved yield in phosphorus limited environments (Beebe et al., 2008). Root traits for improved water uptake include existence of more small fine roots, higher root length as well as higher root density (Fenta et al., 2020; Polania et al., 2017). In drought-tolerant common beans, maturity acceleration with a high seed filling rate further limits any drought impact (Rosales-Serna et al., 2004). Besides improved remobilization and partitioning of photosynthates (Teran et al., 2019), a change of canopy biomass and harvest index has also been found to contribute in common beans to more drought tolerance (Assefa et al., 2015; Polania, Rao, et al., 2016). Hageman et al. (2020) further provided evidence that resource partitioning from pod walls into seeds and the inherent sensitivity of leaflet growth rate to drought can be used as further indicators for drought sensitivity/tolerance. When screening 25 common bean genotypes with contrasting drought tolerance in a phenotyping platform under different water supply conditions, gas exchange and osmotic adjustment together with increase in grain yield were also proposed as useful indicators for selecting more drought-tolerant common bean lines (Lanna et al., 2018). Polania et al. (2016) further reported that the best nitrogen-fixing common bean lines under soil water deficit are more drought-tolerant. But common bean genotypes, even more tolerant to soil water deficit, can ultimately be severely affected by nitrogen-deficiency under such soil water deficit (Beebe et al., 2014).

<figure><img src="image_3.png" /><figcaption>**FIGURE 1** Common bean plants (left) and effect of drought on common beans (right) (adapted from Michigan State University Department of Plant, Soil and Microbial Sciences at https://www.canr.msu.edu/beanbreeding/research/drought)</figcaption></figure>
Abiotic stresses further affect the nutritional quality of common beans (Sica et al., 2021). Variability in rainfall affects, for example, the production of secondary metabolites, such as phenolics. This change directly impacts the bean's health-related benefits and sensory qualities (Ahmed & Stepp, 2016). Mild drought stress increases certain phenolic compounds without affecting the seed yield (Herrera et al., 2019). Studies investigating irrigation effects further found that the total fibre decreases under rain-fed conditions. Water availability, however, also influences the digestibility of bean carbohydrates, extractable polyphenols and, depending on the bean variety, the antioxidant capacity (Ovando-Martínez et al., 2014). Only a few studies have so far investigated in greater detail the impact of stresses associated with climate change, such as drought/heat stress, on the bean's mineral and antinutrient content with phytic acid (PA) a potent antinutrient (Hummel et al., 2018).

The aim of this review is, therefore, to discuss the current existing knowledge on the link between drought/heat stress and content of Fe and PA in common beans. In our review, we will first provide a short general overview on the contribution of common beans to food/nutrient security as well as health. This will include the guidelines dictated by countries to improve human diet-related habits and lifestyle, and the role of antinutrients. We will then review the existing knowledge with regards to Fe and PA content in common beans and how drought/heat stress affects the content of these two compounds. Figure 2 provides a simplified overview of the processes specifically reviewed. A discussion will follow on how breeding using modern “omics” approaches has so far contributed not only to more drought/heat stress tolerance in common beans but also to identify genes involved in drought/heat stress tolerance as well as increase in Fe content and reduction in the PA content. Finally, we will discuss areas for possible future exploration of existing knowledge in common beans regarding increasing Fe and antinutrient content particularly under drought/heat stress conditions.

## 2 | FOOD AND NUTRIENT SECURITY

### 2.1 | Food security

Globally, common beans are grown on 23 million ha (http://www.cgiar.org/our-research/crop-factsheets/beans) and the global common bean production has now risen to 12 million tons (mt) per year (FAO, 2014, 2018; Heinemann et al., 2016). The bean greatly contributes to overall food and nutrient security particularly in sub-Saharan Africa and in Central and Southern American countries (Beebe, 2012; Broughton et al., 2003). Latin America is the region

<figure><img src="image_4.png" /><figcaption>FIGURE 2 Effect of drought/heat stress on minerals and phytic acid in common beans and the consequences of biofortification</figcaption></figure>

with the greatest production of common beans, representing about 50% of world volume, followed by Africa with 25% (Figure 3). In sub-Saharan Africa, common beans are produced on more than 3.5 million ha with production taking place mainly in East Africa, the lakes region and the highlands of Southern Africa, with a combined production of almost 1 mt (Demelash, 2018). In Latin America, per capita annual consumption of common beans ranges from 10–18 kg, whereas in East Africa common annual bean consumption can be as high as 50 kg per capita.

In 2019, Myanmar, India and Brazil were further the top three dry bean producing countries in the world and Myanmar and India produced each over 5 mt (FAOSTAT, 2020; Table 1). In Latin America, Brazil was in 2019 the main producer of dry beans, with about 2.8 mt, followed by Mexico with about 0.9 mt. In Europe, only about 544,330 ha were cultivated with beans, with a production of about 1.9 mt (https://www.pulsesincrease.eu/crops/common-bean; accessed August 2021). Although having the lowest production area of all top dry bean producers, the United States of America (Table 1) achieved the highest bean yield in 2019 (1979 kg ha⁻¹). This was most likely due to a better technological input and also better seed quality. However, in India, despite having the greatest cultivation area, bean yield is still very low (418 kg ha⁻¹; Table 1). Such low yields in countries, such as India, Mexico and Kenya, are very likely due to a low technological input by resource poor farmers, irregularities of rainfall as well as poor seed quality.

Low input agricultural systems further account for the majority of common bean production and small-scale farmers particularly depend on beans for food and income (Kermah et al., 2017). They use, however, poor quality seed material caused by poor storage, seed-borne fungal infection as well as sowing and harvesting under unfavourable environmental conditions (Oshone et al., 2014).
<figure><img src="image_5.png" /><figcaption>FIGURE 3 Impact of climate change on the suitability of bean production (adapted from Beebe et al., 2011). White areas represent areas where common beans are either not extensively grown or where climate change might have very little impact on bean growth</figcaption></figure>

<table><caption>TABLE 1 Top common bean producing countries in the world</caption><thead><tr><th></th><th>Area (ha)</th><th>Production (kg/ha)</th><th>Total production (tons)</th></tr></thead><tbody><tr><td>Myanmar</td><td>3.201.135</td><td>1826</td><td>5.845.272</td></tr><tr><td>India</td><td>12.690.696</td><td>418</td><td>5.309.787</td></tr><tr><td>Brazil</td><td>2.610.585</td><td>1113</td><td>2.897.749</td></tr><tr><td>China</td><td>745.936</td><td>1739</td><td>1.297.182</td></tr><tr><td>United Republic of Tanzania</td><td>893.570</td><td>1340</td><td>1.197.383</td></tr><tr><td>Uganda</td><td>539.660</td><td>1815</td><td>979.482</td></tr><tr><td>United States of America</td><td>470.890</td><td>1979</td><td>931.891</td></tr><tr><td>Mexico</td><td>1.207.395</td><td>728</td><td>878.983</td></tr><tr><td>Kenya</td><td>1.167.543</td><td>639</td><td>746.059</td></tr></tbody></table>

Note: Source: Food and Agriculture Organization Statistical Databases was used to develop this table; FAOSTAT, 2020).

They further grow beans in association, or in rotation, with maize with minimal production inputs (Rurangwa et al., 2018). Cereals are thereby grown on more fertile fields and legumes on soils depleted in nutrients (Chekanai et al., 2018; Kelly et al., 1998). Due to its existing SNF capacity, although lower when compared to soybean and faba beans (Peoples et al., 2009), common beans can grow on such marginal lands. Although common bean varieties with high SNF capacity and environmental resilience would be, therefore, of great benefit (Wilker, 2021), common bean breeding seldom includes selection for the SNF trait. In addition, modern bean production practices involve the application of nitrogen-fertilizer which causes not only SNF downregulation but also environmental pollution (Wilker et al., 2019). To solve this problem, moderate phosphorus fertilization in combination with an appropriate more affordable Rhizobium inoculation as a nitrogen source might be one potential option for

improved production, especially under soil water deficit (Kibido et al., 2020; Samago et al., 2018). Overall, all these problems ultimately limit bean yields to low as ≤0.5 t ha⁻¹. This not only greatly affects smallholder systems (FAO, 2014; Rao et al., 2016), but prevents the realization of the bean's full yield potential and causes production instability from 1 year to the other. Most worrying, based on crop modelling, the majority of current common bean growing areas, especially in south-eastern Africa, will be in 2050 unsuitable for bean cultivation greatly affecting food and nutrient security (Hummel et al., 2018).

As much as one-third of bean production areas are further influenced by drought/heat stress. This greatly affects the contribution of common beans to food and nutrient security (Beebe et al., 2014; Kazai et al., 2019). Common bean production areas particularly subjected to frequent droughts are highland Mexico, the Pacific coast of Central America, northeast Brazil, and eastern and southern
Africa from Ethiopia to South Africa (Beebe et al., 2013). More than 60% of the world's common beans are cultivated under non-irrigated conditions in areas where seasonal rainfall is erratic and beans grow under rain-fed conditions (Seidel et al., 2016; Smith et al., 2019). Drought/heat stress can cause yield losses of up to 80% in these rain-fed production systems (Kazai et al., 2019), Specifically, intermittent or terminal drought stress causes severe yield loss (Beebe et al., 2013), although drought stress towards the end of the growing season might not cause much harm for grain yield (Mathobo et al., 2017). Irrigation would, therefore, allow to increase common bean production. Indeed, in Brazil yields of around 2900 kg ha⁻¹ have been already obtained by irrigation (Alves Souza et al., 2020).

Recent modelling studies raise further concern for future food and nutrient security. They predict a significant decrease in the future suitability to grow common beans due to increased drought and also heat stress (Heinemann et al., 2017). In Africa, for example, where an estimated 682,000 ha of beans are currently cultivated, annual yield loss due to drought/heat conditions is already in the range of 781,000 t. Simulation models to characterize bean production in Brazil in rain-fed environments also indicate that climate change will cause more frequent but less severe drought conditions (Heinemann et al., 2017). A previous ecological diversity study with wild common bean accessions covering a habitat from Mexico to Argentina further found that accessions are distributed among different precipitation regimes following a latitudinal gradient. The habitat ecological diversity of the collection sites was further associated with natural sub-populations (Cortés et al., 2013). Finally, current common bean areas might also shift to colder regions of the Northern Hemisphere, such as Canada, the Nordic countries and Russia, as indicated by recent model projections (Ramirez-Cabral et al., 2016). How these changes will actually affect overall bean production has still to be investigated in more detail.

## 2.2 | Nutrient security

In contrast to the industrialized world, common bean is the most important grain legume consumed in areas with a low income where health is influenced mainly by dietary deficiencies rather than by excesses. Common beans are therefore also regarded, as other pulses, the “poor man’s meat”. As a food staple, common beans contribute up to 35% of the protein and 340 calories/100 grams to the daily diet of resource poor urban and rural families. Food, however, not only needs to satisfy the caloric requirements, but has ultimately to provide sufficient amounts of nutrients such as minerals and vitamins (Muller et al., 2017). Common beans are an important source for the minerals

Fe and Zn (Beebe et al., 2000; Castro-Guerrero et al., 2016; Drewnowski, 2010; Graham et al., 2007; Hall et al., 2017; Mitchell et al., 2009). For a general overview of the nutritional value of common beans, see https://feedtables.com/content/common-bean and also Celmeli et al. (2018). Fe and Zn deficiency further affects over 30% of the world's population (Bailey et al., 2015). Common bean has, therefore, the potential to not only reduce poverty but also to increase nutrient security, particularly on smallholder farms (De Luque & Creamer, 2014). The access to diverse, nutrient-dense food sources is consequently a priority in order to improve sustainable nutrient security, especially in low-income countries and to prevent hidden hunger (Nelson et al., 2018; Petry et al., 2015). Hidden hunger is generally concerned with a deficiency of nutrients and occurs when the food quality in a person's diet is insufficient for normal growth and development. Minerals, such as Fe and Zn, are thereby key determinants in staple crops and foods for sufficient dietary micronutrient uptake (Díaz-Gómez et al., 2017).

Common beans further present a much better source for these minerals in comparison to cereals (Castro-Guerrero et al., 2016). In common beans, the Fe concentration ranges from 35 to 90 µg/g with an average of 55 µg/g and is higher when compared to crops like rice (6.3–24.4 µg/g), wheat (25 µg/g–56 µg/g) or maize (9.6–63.2 µg/g). Breeding approaches have recently even achieved a Fe concentration of 130 µg/g (Kimani & Warsame, 2019). Common beans also have a relatively high Zn seed content (21–54 µg/g), with an average of 35 µg/g. Environmental and genotypes can, however, influence seed mineral concentrations as recently found with landraces and improved common bean varieties (Caproni et al., 2020; Hummel et al., 2018; Murube et al., 2021; Philipo et al., 2020). Although there is no statistically significant correlation between Fe and Zn content of seeds and the geographical distribution of bean (Caproni et al., 2020), previous studies provided evidence for a tendency for Andean pools to contain more Fe (Beebe et al., 2000) and Mesoamerican pools more Zn (Islam et al., 2002). Moreover, in a more recent study, new multi-parent populations were developed at the University of Nairobi. Lines harbouring different tolerance traits (drought, low soil fertility, major bean diseases) were combined with lines with high mineral traits (Fe and Zn). Eighty-four selected lines were more drought-tolerant and had more than 90% better yield compared to their parents. Forty-six promising lines had further higher grain Fe and also Zn concentrations compared with their parents. These lines can possibly now contribute to increased bean productivity and also combating micronutrient deficiencies in eastern Africa and other parts of Africa (Kimani & Warsame, 2019).
3 | HEALTH

3.1 | Impact on health

Pregnant women and children are specifically at risk in their health due to high mineral needs but poor mineral intake. While minerals are required by humans in small quantities, minerals participate in a wide variety of metabolic processes. Fe is, for example, required for the synthesis of haemoglobin and several hormones. A conventional breeding programme developed in Rwanda and in the Democratic Republic of Congo (DRC) already resulted in the selection of high Fe-containing common bean lines. These lines were also well adapted to the local conditions and suited both the farmers' and consumer preferences (Mulambu et al., 2017). The programme HarvestPlus of the Consultative Group for International Agricultural Research (CGIAR) particularly focuses on the selection of such Fe bio-fortified common bean lines (Asare-Marfo et al., 2013). African countries, followed by countries in Latin America, the Caribbean as well as Asia, rank, therefore, high on the list of countries targeted under the Biofortification Priority Index (HarvestPlus, 2020) (Figure 4). Such Fe-biofortified bean lines have already improved the Fe status and health of women in several African countries (Andersson et al.,

2017; Haas et al., 2016). However, how these lines per-
form with regards to yield as well as nutrition under
drought/heat stress conditions associated with climate
change still remains to be investigated.

A previous study on diet modelling also found an association between bean consumption, improvement of nutrient intakes and healthy eating index scores (Hornick, 2007). Low-quality diets often lack dietary diversity along with high amounts of saturated fat and low vegetable, fruit, as well as fibre intake that can contribute to disease risk (Hiza et al., 2013). Consuming dry beans results in higher intake (10% or more) of fibre, protein, folate, Zn, Fe and Mg with lower intake of saturated fat and total fat which provides an improvement in the overall diet quality (Mitchell et al., 2009). Replacing refined carbohydrates in the diet with protein sources that are low in saturated fat, as in beans, reduces, for example, the risk of cardiovascular diseases (Hu, 2005; Mobley et al., 2014). People following a Mediterranean-style diet, richer in plant foods, including bean, have, indeed, a lower risk of cardiovascular disease and mortality (Estruch et al., 2006; Serra-Majem et al., 2006). The Dietary Guidelines for Americans (DGA, 2015–2020) (available at https://health.gov/our-work/food-nutrition/previous-dietary-guidelines/2015, accessed May 2021) further consider the beans' nutritional content

<figure><img src="image_6.png" /><figcaption>FIGURE 4 Common bean Bio-fortification Priority Index. Picture courtesy of HarvestPlus (https://bpi.harvestplus.org/bpi_cropmaps.html?id=c12021)</figcaption></figure>
and benefit as both a protein and a vegetable. Beans can, therefore, be regarded as either in order to appropriately meet the recommended dietary intakes. The Guidelines on Nutrition and Physical Activity for Cancer Prevention advocates the consumption of poultry, fish and beans as an alternative to pork, lamb and beef while stressing the importance of vegetables (Kushi et al., 2012). The American Heart Association further recommends consuming 360 g of beans per week for adults for the prevention of cardiovascular diseases (Van Horn et al., 2016). However, the large consumption of bean seeds, as a “meat replacement” staple food in many countries of Africa, Central and South America and Southern Asia, has already contributed to nutritional problems.

Finally, it should not be overlooked that common bean seeds are also an important protein source (16–30%), despite being deficient in sulphur amino acids (methionine and cysteine). In particular for geographic areas, where a large part of the dietary protein is obtained from legumes, there is a need to increase the content of sulphur containing amino acids in legume proteins. Increasing the concentration of S-containing amino acids is, therefore, a current major research task (Saboori-Robat et al., 2019). Several legume proteins, including those of common bean seeds (e.g. 7S globulins, protease inhibitors, lectins), are now regarded not only as, but also functional ingredients (Carbonaro et al., 2015). Common bean seeds further contain no cholesterol and only low amounts of total and saturated fats. They are rich, however, in carbohydrates (up to 60%), especially starch and several vitamins (biotin, folic acid, niacin, thiamine and riboflavin). They are further an important source of dietary fibre (up to 37%) (Trinidad et al., 2010). Distinctive physiological roles have been attributed to the two different fibre fractions: soluble vs. insoluble. Soluble fibre, making up only a small part of the total dietary fibre, can assist in lowering cholesterol levels as well as decreasing the risk of heart diseases in regulating blood glucose levels. The regulation of intestinal function is affected by the insoluble fibre fraction. Studies have also produced evidence showing the protective effect of legume fibres against risk of developing colon cancer (Zhu et al., 2015, Wang et al., 2013; Campos-Vega et al., 2013). Due to this high fibre content, common beans can significantly contribute to the recommended dietary fibre intake of adult women (25 g/day) and adult men (38 g/day) (Dahl & Stewart, 2015).

## 3.2 | Antinutrients

Common beans are also a source, as other legumes, of bioactive compounds. Bioactive compounds in common bean seeds include oligosaccharides, lectins, phytates,

enzyme inhibitors, phenolic compounds, complex polyphenols (tannins) as well as saponins. The presence of these compounds has promoted the nutraceutical use of legumes (Carbonaro, 2021). In spite of the well-documented antinutritional effects of some legumes, residual (i.e. below their toxic level) amounts of most biologically active compounds help in the prevention and management of severe diseases, such as cardiovascular diseases. These diseases mainly affect the world population of industrialized countries (Padhi & Ramdath, 2017). Hypercholesterolemia (Zhang et al., 2010), type-2 diabetes (Mattei et al., 2015) and cancer (Mitchell et al., 2009) are diseases which can be prevented. Selection to decrease the antinutritional substances have been already done for common bean varieties that are generally consumed. Cooking and processing are two methods that also inactivate trypsin inhibitors that are relevant antinutritional factors. These inhibitors reduce the digestion and absorption of dietary proteins (Avilés-Gaxiola et al., 2018). Removing antinutritional substances may, however, result in yield reductions as they also play an additional protective role in pathogen or insect resistance (Boulter, 1982). Varieties with a high content of heat-stable (non-protein) compounds, such as tannins and PA, pose further concern. This is because protein digestibility is lowered by high molecular weight tannins (Mr 500–5000), especially condensed tannins (proanthocyanidins). They form strong insoluble complexes with proteins. Moreover, the reactivity of some tannins, in particular those in common beans, increases after thermal treatment (Carbonaro et al., 1992). Tannins further adversely affect the absorption of trace elements, especially of Fe, but also of Zn and Cu, as a result of tannin-metal complex insolubilization, particularly after cooking (Carbonaro et al., 2001). As discussed below, it is now also well documented that the other major cation chelator in the seed is PA (reviewed by Petry et al., 2015). Finally, tannins, PA and saponins also interact with the absorption of fat-soluble compounds including fat-soluble vitamins and carotenoids.

## 4 | MINERALS/PA AND DROUGHT/HEAT STRESS

### 4.1 | Mineral/PA content

In order to provide sufficient minerals to millions of consumers, relying on common beans as part of their diet, a better understanding is needed of the effect that particularly drought/heat stress has on bean yield as well as on the bean's mineral and antinutrient content. In this regard, some knowledge already exists from characterizing
lentils. Lentil plants exposed to drought and/or heat stress had lower amounts of vital minerals (K, P, Ca, Fe, Mn and Zn). This decrease was further associated with a reduction of root biomass under heat stress and a negative effect, due to drought stress, on transpiration, stomatal conductance as well as root function. When both stresses (drought and heat) acted simultaneously on lentil plants, the effect was even more profound (Sehgal et al., 2019). Choukri et al. (2020) also analysed 100 lentil genotypes from a global collection grown under normal, heat and combined heat-drought conditions. Fe, Zn and crude protein content was significantly reduced under these stress conditions. The effect of combined heat-drought stress was, however, more severe than by heat stress alone. A significant positive correlation also existed in lentils between Fe and Zn concentrations under both non-stress and stress conditions (Choukri et al., 2020).

Most efforts to improve the nutritional quality of common beans have up to now focused on developing varieties with lower amounts of antinutritional compounds, such as lectins (phyto-hemagglutinin L, phyto-hemagglutinin E), phaseolin and phytates, and a higher Fe and Zn content (Cominelli et al., 2020; Samtiya et al., 2020; Vasconcelos & Oliveira, 2004). For common beans, genotypic variation in the content of Mg, sulphur and Fe in bean leaves, and Ca and Fe in seeds have been reported in response to drought (Beebe et al., 2000). Smith et al. (2019) also found, by analysing a small number of 10 bred lines developed by CIAT, that drought can decrease the amount of minerals in the common bean soluble leaf fraction, but not within the seeds. A reduction in the concentration of Fe as well as phosphorus and Zn by 5-20% have been also reported for a still rather small number of bean lines/varieties under drought conditions (Hummel et al., 2018; Sehgal et al., 2018; Smith et al., 2019). A first multi-year field trial at a climate analogue site experiencing weather conditions, similar which has been predicted in year 2095 for Malawi, has also provided strong evidence that the amount of Fe significantly declines in common beans under drought conditions (Hummel et al., 2018). In contrast, when 20 bean varieties were tested, a significant increase in Zn, due to drought stress, has been measured (Hummel et al., 2018). While changing climatic conditions might, therefore, result in increased Zn-levels in future bean servings, these servings possibly have a lower Fe content and a higher amount of undesirable antinutrients, such as PA (Hummel et al., 2018; Nelson et al., 2018).

PA is the most abundant phosphorylated derivative of myo-inositol and the main storage form of phosphorus in the seed. PA also plays an important function in regulating different cellular processes and also limits oxidative stress (Sparvoli & Cominelli, 2015). The amount of PA, dependent on the phosphate concentration, mainly

accumulates in the bean seed and in the cotyledons (95-98%), with only a small quantity found in the embryo and the seed coat (Blair, Herrera, et al., 2012). PA is further negatively charged and a strong mineral cation chelator. Any increase in the PA amount due to drought/heat conditions is, therefore, of great concern. In experiments carried out in the field with different common bean varieties under rain-fed and drought conditions, representing conditions forecasted by 2050 for south-eastern Africa by the EcoCrop climate impact modelling system, a significant increase in the PA amount in common beans exposed to drought stress was already found. This increase was from 0.96% under rain-fed conditions to 1.16% under drought conditions. Fe, Zn and PA under drought stress conditions at the field site were further influenced by weather conditions rather than genotype (variety) (Hummel et al., 2018). The underlying physiological basis for the PA increase under drought stress is, however, not well studied.

Since data are overall still contradicting and only based on a small number of lines and varieties tested, more in-depth investigations are urgently required to more exactly determine how drought and heat-associated reductions in yield also affects the nutrient quality of common bean seeds and the PA content. Such more detailed investigations should be carried out, however, not only under combined drought and heat stress conditions in a greenhouse but also under natural field conditions to more precisely determine the effect of intense natural drought/heat conditions on minerals and antinutrients.

## 4.2 Mineral uptake and bioavailability

Fe accumulation is highest in leaves with increasing ferritin synthesis during plant development (Zielińska-Dawidziak, 2015). Ferritin functions as the main Fe-storing protein in the seeds of legumes, which has traditionally been the source of plant-derived ferritin (Marentes & Grusak, 1998). Relatively little is, however, currently known about Fe uptake and regulation in legumes, in particular under drought/heat stress conditions. This is despite several articles having recently reviewed Fe uptake and transportation in plants in general (Curie & Mari, 2017; Kobayashi & Nishizawa, 2012; Thomine & Vert, 2013). A large number of likely transport protein candidates have been already identified in legumes as more genome and transcriptome data of various legumes are becoming available. Members of the NRAMP, YSL, VIT and ZIP transport families have higher expression in legume root nodules. These members likely play a role in the transport of Fe across symbiotic membranes (Brear et al., 2013). The application of fertilizers, either to the soil or as a foliar spray, also significantly increases the accumulation
of nutrients in the seed, but transport is hampered by lim-
ited mobility in the phloem sap (Bindraban et al., 2015).

Nutrients can be further relatively immobile, not only
in plant tissues, but also in the soil. This immobility af-
fects their transport within the phloem. A greater rooting
depth is generally required for plants to access from the
soil sufficient amounts of minerals (Maillard et al., 2015).
Transporters, which are essential for the uptake of miner-
als from the rhizosphere, have been characterized in com-
mon beans (Castro-Guerrero et al., 2016). Fe and ferritin
further accumulate in separate cellular locations in bean
seeds. Fe primarily accumulates in the cytoplasm of cells
surrounding the pro-vascular tissue, while ferritin, the
major Fe-storing protein in legume seeds, accumulates in
the amyloplast, as found for peas (Cvitanich et al., 2010;
Marentes & Grusak, 1998). In common bean seeds, only
15% to 30% of total Fe is, however, ferritin bound. An ex-
cess of Fe and osmotic stress increases ferritin expression
in common beans (Hoppler et al., 2014), but ferritin does,
overall, not contribute much in the provision of Fe.

IRT-like transporters are further involved in the uptake
of both minerals that are then mobilized to the shoots via
the xylem and then delivered to developing tissues, includ-
ing seeds, exclusively via the phloem (Hindt & Guerinot,
2012; Khan et al., 2014; Sinclair & Krämer, 2012). Fe up-
take regulation is under the control of two transcriptional
networks, FIT (At2g28160) and PYE (At3g47640), while
Zn uptake requires bZIP transcription factors (Hindt &
Guerinot, 2012; Sinclair & Krämer, 2012). The common
bean genome contains putative homologs for the com-
ponents of the networks (Phvul.005G130500/FIT1-like;
Phvul.002G099700/IRT1-like; Phvul.003G086500/OPT3-
like; Phvul.011G035700/bZIP23-like) (Castro-Guerrero
et al., 2016). Furthermore, an Arabidopsis protein local-
ized in the phloem, OPT3, is a component of the shoot-to-
root signalling network. This protein, not yet characterized
in more detail in common beans, passes on the Fe status in
leaves to roots and *opt3* mutants accumulate more Fe and
Zn in roots and leaves (Mendoza-Cózatl et al., 2014).

The presence of antinutritional compounds, such as
PA and polyphenols, limits the bioavailability of min-
erals (Glahn et al., 2017; Petry et al., 2010; Tako et al.,
2014). By adversely affecting the absorption of minerals,
PA decreases the bioavailability of these minerals and,
as a consequence, negatively impacts the nutritional
value of seeds (Petry et al., 2015). The amount of PA
has been further positively correlated with amounts of
non-ferritin bound Fe (DeLaat et al., 2014). Polyphenols
are also highly interconnected with mineral amounts
(e.g. Fe). They are further involved in resistance to dif-
ferent types of stresses, in part due to their antioxidant
properties (Herrera et al., 2019). A major obstacle to

Fe biofortification in common beans is, therefore, low Fe absorption due to the presence of polyphenol(s) (Ganesan & Xu, 2017). Different subclasses of such polyphenols are present in common beans, mainly located in the seed coat. Although some polyphenols, such as kaempferol, kaempferol 3-glucoside, catechin and 3,4-dihydroxybenzoic acid, are able to promote Fe uptake, at least in an *in vitro* system, others have a strong inhibitory effect. This outweighs the effect of those compounds promoting an increase in Fe uptake (de Figueiredo et al., 2017; Hart et al., 2015). Unfortunately, to our knowledge, a serious gap is still the lack of any bean breeding programme specifically targeting polyphenols for reduction of any negative effects these polyphenols have particularly on Fe bioavailability and how drought/heat stress can affect this process.

5 | BREEDING USING “OMICS”
TOOLS

Previous common bean research work, particularly on drought/heat tolerance, focused mainly on investigating and characterizing agro-morphological traits to identify bean lines with better plant growth under these stresses. Several different traits, for example pod harvest index, were thereby found to be associated with drought tolerance (Polania, Rao, et al., 2016). “Omics” tools (genomics, transcriptomics and proteomics) have been, or are currently, applied to identify and characterize genes and genome sequences in common bean plants that are involved in drought/heat stress tolerance or mineral production. These activities are aimed to produce improved common bean material for drought/stress tolerance and higher mineral content. A recent review has highlighted the achievements in common beans by applying “omics” tools (Nadeem et al., 2021).

Most of the recent/current advances made by applying
“omics” tools were/are only achievable due to the publi-
cation of the common bean genome sequence (Schmutz
et al., 2014; Vlasova et al., 2016). In contrast to the Andean
common bean genome with an estimated size of 587 mega
base pairs (Mbp) with ~27 thousand genes (91% clustered
in synteny blocks with Glycine max), the Mesoamerican
genome is 549.6 mega base pairs in size with ~30 thousand
genes and 94% of which has been functionally annotated
(Vlasova et al., 2016). The availability of the common bean
sequence in 2014 has further offered the opportunity for
better understanding drought adaptation and tolerance in
common beans. However, previous studies never focused
on a deeper level on any possible link between mineral
accumulation and availability and drought/heat stress.
# 5.1 | Quantitative trait loci (QTL) identification

## 5.1.1 | QTLs for drought tolerance

Advances in genetic investigation using genomics paved in common beans the way not only to better understand genetic variation, for example within the European common bean germplasm and to trace its divergence from the American germplasm (Bellucci et al., 2021; Caproni et al., 2020; Lioi & Piergiovanni, 2013) but also for the identification and characterization of quantitative trait loci (QTL) related to agronomic and nutrient traits. QTL analysis generally links phenotypic and genotypic data. In particular, numerous QTL studies in common beans have been carried out to identify specific traits linking abiotic stresses—including drought—to agronomic traits of interest such as plant size, seed yield and flowering time. These approaches rely on the genetic analyses on crosses of susceptible x tolerant parental common bean genotypes, belonging to either a single gene pool, or both the Andean and Mesoamerican gene pools (Nadeem et al., 2021). The resulting population is then used to construct genetic linkage maps. The resolution of these maps has considerably improved over the years. Based on these maps, an increasing amount of genetic markers including microsatellites and single-nucleotide polymorphisms (SNPs), a variation at a single position in a DNA sequence, have been recently developed (Leitão et al., 2021; Sedlar, Kidrič, et al., 2020; Sedlar, Zupin, et al., 2020). Particularly, mining in common beans QTLs affecting field performance and nutrient value under drought will be still crucial in the future design of molecular tools for marker-associated selection (MAS). Mining will also be crucial for the identification of possible molecular targets important for gene editing (GE) approaches, as recently described for wheat grown in dry and hot environments (Tura et al., 2020). QTLs related to yield, flowering time as well as days to maturity, were specifically identified in common beans based on the genetic analysis of a Mesoamerican x Mesoamerican drought-tolerant x susceptible cross (Blair, Galeano, et al., 2012). In addition, QTLs related to seed yield and flowering time have been recently identified after crossing two Andean genotypes reacting differently to drought stress (Dramadri et al., 2019). Three loci related to seed yield per plant (Syp1.1, Syp1.2, Syp2.1) are of specific interest for yield-oriented MAS under drought stress with Syp1.1 emerging as a master regulator of yield under drought (Sedlar, Zupin, et al., 2020; Trapp et al., 2015). Moreover, a great number (189) of QTLs have been found related to seed weight and 33 QTLs related to yield. Many of these QTLs are within—or in proximity—to genes known to be involved in primary or specialized metabolism

(Valdisser et al., 2020). Finally, a stable QTL related to yield (Yd4.1) has been recently identified in a common bean BAT881 × G21212 RIL population tested in field trials across four different locations in Colombia. This QTL is not only associated with drought stress, but also to phosphorus and Al stress. The molecular function of Yd4.1 is, however, still unknown (Diaz et al., 2018).

Further, SNPs determination in common beans allowed to annotate a SNP to a gene related to drought tolerance. This SNP is related to biosynthesis of proline, a well-known osmotic protector (Villordo-Pineda et al., 2015). A most recent SNP analysis under drought conditions also identified SNPs for processes responsive to drought stress. These processes included stomatal regulation, protein translocation across membranes, redox mechanisms, hormone as well as osmotic stress signalling (Leitão et al., 2021). A further more recent whole-genome resequencing-derived SNP dataset applied for a genome-wide association analysis identified 12 loci. These loci were significantly associated with survival after drought stress at the seedling stage. They also confirmed the drought-related function of an aquaporin gene (PvXIP1;2) located at Locus_10 (Wu et al., 2021). Finally, a good example of what can be achieved by applying the QTL technology in legumes has been recently shown for chickpea and cowpea. Genetic physical maps were developed and QTLs including “QTL-hotspot” regions containing QTLs for several drought tolerance traits were identified. This analysis has already resulted in 29 new cowpea varieties developed and the production of 20,353 t of certified seeds which have been planted on about 508,825 ha (Varshney et al., 2019).

## 5.1.2 | QTLs for mineral (Fe) content

Studies about the genetic basis of common bean seed composition have so far mainly focused on minerals, such as phosphorus, Fe and Zn, due to their problems related to deficiency in human diets. Identification of QTLs was thereby based on the application of both inter and intragene pool populations (Casañas et al., 2013). A recent genome-wide association studies (GWAS) resulted in the identification of quantitative trait nucleotides (QTNs) associated with seed content of nitrogen, phosphorus, Ca, Mn and Zn, while no significant associations were found for Fe content (Gunjača et al., 2021). In contrast, Blair, Galeano, et al. (2012) found numerous QTLs, also related to Fe, although usually found to be population or environment specific. QTLs associated to seed phosphate content have been identified in a RIL common bean population (intragene pool Andean x Andean) on chromosomes Pv02, Pv05, Pv06, Pv05 and Pv11, with additional PA-related
QTLs on Pv04 and Pv08 (Cichy et al., 2009). P-QTLs derived from the analysis of intergene pool populations (Mesoamerican × Andean). Loci associated to seed phosphate and total phytates were identified on Pv02, Pv03, Pv04, Pv06, Pv10 and Pv11 (Blair, Galeano, et al., 2012). An intergene pool study based on a Mesoamerican × Andean cross further identified QTLs associated with both Fe and Zn content. QTLs were scattered along chromosomes Pv03, Pv04, Pv06, Pv07, Pv08, Pv09, with a cluster of 5 on Pv11 (Blair et al., 2010). Furthermore, overlapping Fe and Zn-QTLs were identified on a linkage group located on Pv06, alongside QTLs located on Pv03, Pv04, Pv07, Pv08 and Pv11 (Blair et al., 2010). A recent meta-analysis, conducted on the cited literature, finally reduced the original set of detected QTLs into a set of 12 QTLs, with two QTLs specific for Fe and Zn, and 8 QTLs related to both (Izquierdo et al., 2018).

Interestingly, a recent study has presented the first common bean MAGIC population of the Mesoamerican gene pool (Diaz et al., 2020). The study allowed the identification of different genomic regions associated with yield, mineral accumulation, phenology and physiological traits under drought conditions. Moreover, major QTLs controlling more than one trait, even in different seasons, and candidate genes for major QTLs were identified. This study now provides interesting data for the development of advanced breeding tools. In a further recent development, optimal contributions selection was applied to design common bean crossings within four market groups with relevance for East Africa. Genomic estimated breeding values were thereby predicted for grain yield, cooking time, Fe, and Zn in an African bean panel of 358 genotypes in a two-stage analysis. Such genomic selection using optimal contributions selection will possibly accelerate breeding of high-yielding, biofortified, and rapid cooking African common bean cultivars (Saradadevi et al., 2021).

## 5.2 | Biotechnology

"Biotechnology" to improve common beans was already suggested in 2003 (Svetleva et al., 2003). Plant biotechnology generally allows precise genetic changes by integrating, for example, an identified and characterized gene providing a beneficial trait into the plant genome or to change a gene inside a plant by genome editing (Du et al., 2016). The process to obtain either a transgenic genetically modified plant (GMO) or gene-edited plants generally involves as tools application of plant transformation to insert a gene sequence, in vitro culture of transformed plant tissues as well as whole plant regeneration. Although potential genes for transformation are

known, due to an available sequenced common bean genome with 94% of genes functionally annotated (Vlasova et al., 2016) allowing to identify target genes for any desirable trait modification, efficient bean transformation to obtain transformed modified plants has still technical limits. Common bean transformation is by far not a routine approach, as in other species (De Paolis et al., 2019). However, first examples of possible successful genetic modification of common beans include overexpression of a methionine-rich storage albumin from Brazil nut in transformed bean plants after particle bombardment of the apical meristematic region of embryos for gene transfer (Aragão et al., 1999). Expression of the barley (Hordeum vulgare) late embryogenesis abundant protein (HVA1) in transformed common beans is a further example where the method of particle bombardment of the shoot meristem for transformation was applied (Kwapata et al., 2012). Produced transformed plants were more drought-tolerant due to longer roots. These examples provide overall evidence that plant transformation is, indeed, applicable for bean biofortification.

### 5.2.1 | Genes and drought/heat stress tolerance

Blair et al. (2011) already characterized 4219 uniGENES from cDNA libraries prepared from contrasting drought-treated common bean genotypes. By applying suppression subtractive hybridization (SSH) and a whole-genome protein database for target hits, TRANSCRIPTION FACTORS (NAC and AP2-EREBP family) and genes involved cell metabolic processes and present in photosynthesis were further identified to be involved in the drought response of common bean (Müller et al., 2014; Recchia et al., 2013; Wu et al., 2016). Table 2 shows a selection of genes which have been so far investigated regarding drought stress in common beans. When further drought responsive genes in leaf and root tissue of common bean were investigated by RNA-Seq, genes were predominantly involved in oxidative stress. This suggests a tolerance mechanism based on reduction of damage from reactive oxygen species (Pereira et al., 2020). Orthologues of the soybean Hsp20 genes are further up-regulated in response to drought and salinity stress (Lopes-Caitar et al., 2013). López-Hernández and Cortés (2019) recently identified in common beans by coupling genome-environment associations with last-generation genome wide association study algorithms candidate genes including HSP20, but also MED23, MED25, HSFB1 and HSP40 that are directly linked to heat-responsive pathways. Additional candidate genes
<table><caption>TABLE 2 Stress responsive genes and their function in common beans</caption><thead><tr><th>Study</th><th>Function</th><th>Description</th><th>Dry bean accession</th><th>Homologue accession/ species</th></tr></thead><tbody><tr><td rowspan="15">Transcriptome analysis of differentially expressed genes in roots of BAT 447 under drought stress during development (Recchia et al 2013)</td><td rowspan="2">NAC transcription factor (TF)</td><td>NAC domain protein, IPR003441</td><td>|75749297|</td><td>|224088037| Populus trichocarpa</td></tr><tr><td>NAC4 protein</td><td>|75748424|</td><td>|62546189| Glycine max</td></tr><tr><td rowspan="2">DREB TF</td><td>NAC domain protein</td><td>|75749318|</td><td>|224088037| P. trichocarpa</td></tr><tr><td>NAC domain protein</td><td>|75748418|</td><td>|187940303| G. max</td></tr><tr><td rowspan="2">ERF TF</td><td>Fe-S cluster assembly protein DRE2 homolog</td><td>|75749717|</td><td>|292630743| G. max</td></tr><tr><td>DREB</td><td>|75748469|</td><td>|32480821| G. max</td></tr><tr><td rowspan="2">bHLH TF</td><td>Ethylene-responsive element binding factor 4</td><td>|75749028|</td><td>|190361165| G. max</td></tr><tr><td>Transcription factor EIL2</td><td>|75749407|</td><td>|18643339| Vigna radiate</td></tr><tr><td rowspan="2">bZIP TF</td><td>Coiled-coil-helix-coiled-coil-helix domain containing protein</td><td>|75749257|</td><td>|66947630| Medicago truncatula</td></tr><tr><td>Transcription factor bZIP70</td><td>|75749123|</td><td>|145652341| G. max</td></tr><tr><td rowspan="4">MYB</td><td>Leucine-rich repeat protein</td><td>|75748580|</td><td>|223452524| G. max</td></tr><tr><td>TGA-type basic leucine zipper protein</td><td>|75748298|</td><td>|15148922| Phaseolus vulgaris</td></tr><tr><td>F-box/LRR-repeat protein, putative</td><td>|75748883|</td><td>|255558466| Ricinus communis</td></tr><tr><td>MYB transcription factor MYB185</td><td>|75748729|</td><td>|110931684| G. max</td></tr><tr><td>GATA-factors</td><td>GATA transcription factor, putative</td><td>|75748743|</td><td>|255572876| R. communis</td></tr><tr><td>WRKY family</td><td>WRKY36</td><td>|75748775|</td><td>|151934195| G. max</td></tr><tr><td>Ubiquitous factors TFIIA e Sp1</td><td>Transcription initiation factor ia, putative</td><td>|75748702|</td><td>|255566898| R. communis</td></tr><tr><td rowspan="2">IAA (auxin-responsive)</td><td>Auxin-responsive protein IAA1, putative</td><td>|75748737|</td><td>|255552973| R. communis</td></tr><tr><td>Auxin-responsive family protein</td><td>|75748789|</td><td>|15226425| Arabidopsis thaliana</td></tr><tr><td rowspan="2">GRAS</td><td>GRAS family transcription factor</td><td>|75748648|</td><td>|224106445| P. trichocarpa</td></tr><tr><td>GRAS family transcription factor</td><td>|75749650|</td><td>|224106445| P. trichocarpa</td></tr><tr><td>Heteromeric factors</td><td>Transcription factor CCAAT</td><td>|75748712|</td><td>|193237557| Lotus japonicas</td></tr><tr><td rowspan="2">eIF2—alpha family</td><td>Translation initiation factor EIF-2b</td><td>|75748325|</td><td>|255544025| R. communis</td></tr><tr><td>Eukaryotic translation initiation factor 5A</td><td>|75748617|</td><td>|20138704| Manihot esculenta</td></tr><tr><td>Zinc finger</td><td>C2-H2 zinc finger protein</td><td>|75749674|</td><td>|161087182| G. max</td></tr></tbody></table>

(Continues)
<table><caption>TABLE 2 (Continued)</caption><thead><tr><th>Study</th><th>Function</th><th>Description</th><th>Dry bean accession</th><th>Homologue accession/ species</th></tr></thead><tbody><tr><td rowspan="6">Transcriptome analysis of differentially expressed genes in BAT 447 under drought stress during flowering (Müller et al. 2014)</td><td></td><td>Uncharacterized protein LOC100305788</td><td></td><td>|351721030| G. max</td></tr><tr><td></td><td>Oxygen-evolving enhancer protein</td><td></td><td>|358344003| M. truncatula</td></tr><tr><td></td><td>Chlorophyll a/b binding protein type II</td><td></td><td>|16805332| G. max</td></tr><tr><td></td><td>Hypersensitive induced reaction protein 1</td><td></td><td>|354683205| G. max</td></tr><tr><td></td><td>Invertase/pectin methylesterase inhibitor family protein</td><td></td><td>|297310623| Arabidopsis lyrata</td></tr><tr><td></td><td>Auxin-repressed protein</td><td></td><td>|357446689| M. truncatula</td></tr><tr><td rowspan="7">Transcriptome analysis of differentially expressed genes in BAT 447 under drought stress during grain filling (Müller et al. 2014)</td><td></td><td>Predicted: 40S ribosomal protein S17-4</td><td></td><td>|356521554| G. max</td></tr><tr><td></td><td>Predicted: 40S ribosomal protein S16-like</td><td></td><td>|356524632| G. max</td></tr><tr><td></td><td>Leucine zipper protein</td><td></td><td>|357491217| M. truncatula</td></tr><tr><td></td><td>Unknown</td><td></td><td>|388517649| L. japonicus</td></tr><tr><td></td><td>Predicted: cell wall/vacuolar inhibitor of fructosidase 1-like</td><td></td><td>|502150782|Cicer arietinum</td></tr><tr><td></td><td>NAD-dependent isocitrate dehydrogenase</td><td></td><td>|3790188| Nicotiana tabacum</td></tr><tr><td></td><td>Predicted: RING-H2 finger protein ATL66-like</td><td></td><td>|356539989| G. max</td></tr><tr><td rowspan="10">Transcriptional analysis of drought induced genes in the roots of BAT 477 (Recchia et al. 2013)</td><td rowspan="5">Cellular<br>metabolism</td><td rowspan="5">Pyruvate decarboxylase, putative</td><td></td><td>|255579310| R. communis</td></tr><tr><td>Malate dehydrogenase-like protein</td><td></td><td>|83283965| Solanum tuberosum</td></tr><tr><td>Glyceraldehyde-3-phosphate dehydrogenase</td><td></td><td>|255638912| G. max</td></tr><tr><td>Glutaredoxin-1, grx1, putative</td><td></td><td>|255540625| R. communis</td></tr><tr><td>Spliceosomal complex</td><td></td><td>|224094081| P. trichocarpa</td></tr><tr><td rowspan="4">Biological<br>processes</td><td>Methionine adenosyltransferase</td><td></td><td>|75304713| Phaseolus lunatus</td></tr><tr><td>S-adenosylmethionine decarboxylase</td><td></td><td>|156181612| P. vulgaris</td></tr><tr><td>Methionine adenosyltransferase</td><td></td><td>|75304713| P. lunatus</td></tr><tr><td>Interferon-related developmental regulator family protein</td><td></td><td>|42571665| A. thaliana</td></tr><tr><td rowspan="2">Abiotic stress response</td><td>Light-inducible protein ATLS1,</td><td></td><td>|192910730| Elaeis guineensis</td></tr></tbody></table>
<table><caption>TABLE 2 (Continued)</caption><thead><tr><th>Study</th><th>Function</th><th>Description</th><th>Dry bean accession</th><th>Homologue accession/ species</th></tr></thead><tbody><tr><td rowspan="11"></td><td rowspan="11">Biotic stress response</td><td>Group 3 late embryogenesis abundant protein</td><td></td><td>|75708857| P. vulgaris</td></tr><tr><td>Proline-rich protein</td><td></td><td>|806310| G. max</td></tr><tr><td>LEA5</td><td></td><td>|1732556| G. max</td></tr><tr><td>LEA protein</td><td></td><td>|1350522| Picea glauca</td></tr><tr><td>LEA5</td><td></td><td>|1732556| G. max</td></tr><tr><td>Isoflavone synthase 1</td><td></td><td>|184202203| Vigna unguiculata</td></tr><tr><td>Isoflavone synthase 1</td><td></td><td>|184202203| V. unguiculata</td></tr><tr><td>PvPR2</td><td></td><td>|130835| P. vulgaris</td></tr><tr><td>Plastidic phosphate translocator-like protein1</td><td></td><td>|61651606| Mesembryanthemum crystallinum</td></tr><tr><td>Cation:cation antiporter</td><td></td><td>|255587991| R. communis</td></tr><tr><td>ATP binding protein,</td><td></td><td>|255552798| R. communis</td></tr><tr><td>Calcium ion binding</td><td></td><td>|255637247| G. max</td></tr><tr><td rowspan="16">Transcriptional response to drought stress in roots and leaves of drought-susceptible and drought tolerant common bean genotypes (Pereira et al. 2020)</td><td>Aquaporin NIP</td><td></td><td>Phvul.006G171000</td></tr><tr><td>Peripheral-type benzodiazepine receptor and related proteins</td><td></td><td>Phvul.001G205900</td></tr><tr><td>DNAj homolog subfamily c member</td><td></td><td>Phvul.006G060700</td></tr><tr><td>Beta-fructofuranosidase</td><td></td><td>Phvul.005G158500</td></tr><tr><td>Class IV chitinase, insoluble isoenzyme WINV1-related</td><td></td><td>Phvul.005G155800</td></tr><tr><td>Protein phosphatase 2C</td><td></td><td>Phvul.001G075400</td></tr><tr><td>Glutathione S-transferase</td><td></td><td>Phvul.008G113700</td></tr><tr><td>Heat shock transcription factor</td><td></td><td>Phvul.007G061800</td></tr><tr><td>Late embryogenesis abundant (LEA) group 1</td><td></td><td>Phvul.007G259400</td></tr><tr><td>Linoleate 13S-lipoxygenase.</td><td></td><td>Phvul.002G228700</td></tr><tr><td>MYB-like DNA-binding domain</td><td></td><td>Phvul.002G184600</td></tr><tr><td>No apical meristem (NAM) protein</td><td></td><td>Phvul.005G084500</td></tr><tr><td>NADH oxidoreductase-related</td><td></td><td>Phvul.003G131500</td></tr><tr><td>Peroxidase</td><td></td><td>Phvul.009G140700</td></tr><tr><td>Glycosyl hydrolase family 10</td><td></td><td>Phvul.009G120500</td></tr></tbody></table>

involved in the response of common bean to water deficit (drought) conditions were very recently identified from a collection of more than 150 Portuguese common bean accessions (Leitão et al., 2021).

### 5.2.2 | Genes and mineral (Fe) content

“Omics” tools have also been applied in order to increase the mineral content and bioavailability of common
beans. The first important step in lowering the pro-
duction of PA in common beans has been the isolation
and sequencing of genes involved in PA biosynthesis
and transport (Fileppi et al., 2010; Panzeri et al., 2011).
Recently, additional putative biosynthesis and transport
genes have been further identified (Cominelli et al., 2017;
Cominelli, Pilu, et al., 2020). The availability of this data
will now allow new cutting-edge innovative research,
including epi-genomics and translatome analysis. Two
identified allelic common bean mutants, affecting the
PvMRP1 PA transporter, caused a 75–90% reduction in
the PA content (Campion et al., 2009; Cominelli et al.,
2018; Panzeri et al., 2011). Particularly, the mutant bean
line, *lpa1*, has a 25% reduction in raffinosaccharides, the
sugars causing flatulence. The biosynthesis of these sug-
ars is strictly linked to the biosynthesis of PA. A study
with human volunteers further found that seeds from
the *lpa1* mutant line provides better Fe absorption, com-
pared to a non-mutant line (Petry et al., 2014). When ap-
plied in common household recipes, *lpa1* mutant seeds
had, however, a lower retention of Zn. Due to a hard-
to-cook phenotype, associated with the increased ther-
mal stability of lectins in the *lpa1* mutant lines, adverse
gastrointestinal symptoms occurred (Petry et al., 2016).
The effect of the *lpa1* mutation on thermal stability of
seed lectins is, however, only problematic in a genetic
background which contains phyto-hemagglutinin L. In
contrast, no significant effect on thermal stability has
been found when the genetic background contains both
phyto-hemagglutinin-L and phyto-hemagglutinin E,
which most common bean genotypes have (Cominelli,
Galimberti, et al., 2020; Cominelli, Pilu, et al., 2020).

When developing *lpa* mutants, the essential role of
PA as regulator of cellular processes in plant vegetative
tissues has to be considered in order to avoid important
undesirable pleiotropic effects (Sparvoli & Cominelli,
2015). Importantly, the common bean *lpa1* mutant has
no reduced germination or any reduced plant growth
and fertility. Still lacking is, however, a much more de-
tailed morphological/physiological evaluation of such
common bean *lpa1* mutants. Cereal *lpa* mutants, the
first *lpa* mutants isolated, have received so far very little
interest. They are affected in the transporter orthologues
of PvMRP1 (Colombo et al., 2020; Sparvoli & Cominelli,
2015). These cereal mutants have further a reduced yield
(5–10% decrease) and non-optimal field performance
(Raboy, 2020). In contrast, the *lpa1* common bean mu-
tant has no such negative agronomic effects under field
conditions (Campion et al., 2009; Chiozzotto et al.,
2018). However, field studies are urgently required to
assess the potential of this mutant particularly under
field conditions in much more depth. They would be
also interesting for breeding programmes aimed to

develop beans with increased mineral bioavailability
and mineral content as well as lower concentrations of
certain polyphenolic compounds (Hummel et al., 2020).
Molecular markers for the *lpa1* and also *lpa12* bean mu-
tants have been already developed. Such markers can
now be applied in marker-assisted selection of common
bean breeding lines as well as the evaluation of the per-
formance of such lines with either individual or com-
bined traits (Cominelli et al., 2018; Panzeri et al., 2011).
However, some concern with the use of *lpa* beans still
exists. PA, as a broad-spectrum antineoplastic agent, can
act in cancer development and progression (Vucenik,
2019), despite the fact that no phytate has been detected
in human biofluids (Wilson et al., 2015). Consequently,
*lpa* beans may be particularly useful in areas where mi-
cronutrient deficiencies are prevalent. In contrast, crops
with high amounts of phytates can also be beneficial for
health in societies that have in their diet sufficient Fe
available, but where both obesity and cancer are on the
rise (Blair, 2013).

The exact role and function of PA in drought/heat
tolerance is, however, still unclear. So far only known
is that some low phytic acid (*lpa*) mutants are more
drought sensitive (Cerino Badone et al., 2012). An
Arabidopsis thaliana *mrp5* mutant (an *lpa* mutant; Nagy
et al., 2009) and common bean *lpa1* mutants, affected in
the AtMRP5 and PvMRP1 orthologous genes, have so far
found to have some better drought tolerance (Chiozzotto
et al., 2018; Colombo et al., 2020; Klein et al., 2003).
Arabidopsis *mrp5-1* mutant rosette leaves have, in this
regard, closer stomata to prevent water loss, and have
a reduced transpiration rate and improved water use
efficiency (Klein et al., 2003). More drought tolerance
of the common bean *lpa1* mutant is also evident under
symbiosis. Transcriptional data provide evidence of
higher expression of stress-related genes in the nodules
and bacteroids of *lpa1* mutants when compared to nod-
ules from non-mutant plants (Chiozzotto et al., 2018).

Finally, results with the *lpa1* mutant plants now open
a new perspective in obtaining mineral improved com-
mon bean varieties. These varieties should not only bet-
ter cope with drought/heat stress but also provide beans
with a low PA content (Raboy, 2020). Introgression of
these mutations into cultivated bean varieties is, conse-
quently, a current major research task (Campion et al.,
2009; Cominelli et al., 2018; Mulambu et al., 2017). This
task also includes the evaluation of the performance of
such lines, with either individual or combined traits,
under environmental stress conditions. Common beans
cooking and nutritional properties will also be evalu-
ated in more depth in a quest to develop bio-fortified
common bean lines devoid of negative traits (Cominelli,
Galimberti, et al., 2020).
### 5.2.3 | Gene editing

Genome editing (GE) applying the clustered regulatory interspaced short palindromic repeats editing (CRISPR)/Cas system have already been applied to edit certain target genes in legumes (Bhowmik et al., 2021; Ji et al., 2019; Liu et al., 2019). GE is currently an effective “omics”-based tool in the manipulation of traits in crops (Du et al., 2016; Tiwari et al., 2020). While CRISPR is usually considered as a tool to generate double strand breaks, and consequently generate knock-out mutations, the modular nature of the CRISPR technology allows alteration of transcriptional activity, or epigenetic status, at a chosen target site (Lee et al., 2019). This can be achieved with a nuclease-deficient version system (dCas9), which can be tied to a diverse array of epigenetic effector domains for site-specific epigenetic modifications (Pulecio et al., 2017). The current advancements and limitations of GE, particularly in orphan crops, have been recently discussed by Venezia and Creasey Krainer (2021). Innovative techniques, such as GE and speed breeding, might effectively also shorten the time to develop drought-resilient common beans and consequently limit any risk of global food insecurity (Bhowmik et al., 2021). In particular, the development of alternative *lpa* common bean mutant lines by applying GE could prove valuable in the pursuit of improving specifically the nutritional value of common beans. Mutations in the rice OsSULTR3;3 gene, encoding type-3 sulphate transporters, have already resulted in rice *lpa* mutants that, in addition to reduced levels of PA, had changes in the amount of a broad spectrum of compounds such as amino acids, organic acids (e.g. citric acid) and other nutritionally relevant compounds including γ-aminobutyric acid (Zhou et al., 2018).

## 6 | AREAS FOR FUTURE EXPLORATION

To achieve the overall goal of higher common bean yield and dietary quality under changing climatic conditions will certainly require the establishment and application of an integrated research framework. This framework should consist of genomics, systems biology, physiology, as well as modelling and breeding (Palit et al., 2020). Recent advances in sequencing and phenotyping methodologies, the rapidly emerging genetic and genomic resources as well as integrated crop modelling and predictions of climate impacts, supports the establishment of a framework also for common beans. Exploring in more depth how landraces cope with drought/heat stress and why they seemingly have a higher mineral content (especially Fe and Zn) and

protein than modern varieties (Celmeli et al., 2018) and how to produce more efficiently GMOs should also be part of the research activities. In addition, production of more annotated genomes will be very helpful to support any future transcriptomic, proteomic as well as epi-genomic data-mining efforts (Li et al., 2017). Finally, exploring the interesting idea of common bean rewilding, which is the reintroduction of specific traits from wild lines into the genetic background of commercial cultivars, should be part of the activities. Although whole-genome sequence data exist for numerous legume species, including common bean, next-generation sequencing (NGS)-driven improvements have not kept pace with that of cereal crops (Rehman et al., 2019). NGS would, for example, support the rewilding idea. Rewilding will specifically address the loss of diversity during the bean's domestication process and will possibly allow improving the bean's nutritional value and tolerance to stresses (Cowling et al., 2015).

### 6.1 | Exploring drought/heat stress tolerance

For most grain legumes, breeders mainly investigated in the past consequences of drought/heat stress on above-ground traits. However, investigating the relationship between below and above-ground traits by studying in the future in more depth will be an important aspect (Sofi et al., 2021). The application, specifically of proteomics as an “omics tool”, will particularly allow to explore proteins involved in drought/heat tolerance and mineral production. Such proteomics studies will allow us to also investigate how these proteins are regulated (Zargar et al., 2017). Furthermore, specific target genes require more in depth investigation. Genes include the orthologous forms of the *Arabidopsis thaliana* and *Vitis vinifera* MYB60 genes. These genes, not well characterized in common beans, have been already extensively characterized for their specific role in the modulation of stomatal movement (Cominelli et al., 2005; Galbiati et al., 2011). Other possible target genes to explore are regulatory genes controlling the expression of DREB genes and that are activated by drought stress (Marcolino-Gomes et al., 2014).

### 6.2 | Improving mineral availability

An important future research priority is increasing mineral (Fe)-bioavailability in common beans. Fe biofortification in common beans requires, however, adequate Fe partitioning between plant tissues. The Fe, and also Zn, uptake mechanism as well as mobilization to allocate more Fe and Zn into bean seeds is, therefore, an area to
be explored in more depth, particularly under drought/ heat stress conditions. Specifically, IRT-like transport- ers involved in the mineral uptake can be thereby in- vestigated. Since these transporters are also under the control of two transcriptional networks, these networks can be specifically characterized for how drought/heat stress affects these networks. Furthermore, isolation and characterization of the protein OPT3, a component of the shoot-to-root signalling network passing on the Fe status in leaves to roots, is a worthwhile target for more in depth investigation.

Different possible bio-fortification strategies to increase amounts of minerals have been already reviewed by White and Broadley (2009). Fe biofortication might also include the future isolation of common bean genotypes low in PA content. To our knowledge, no extensive variability for the PA trait, as found in mung beans (Dhole & Reddy, 2015), has been so far described for common beans. Only two *lpa* mutants have been isolated and characterized in common beans (Campion et al., 2009; Chiozzotto et al., 2018; Cominelli et al., 2018; Panzeri et al., 2011). Selection of more low PA bean mutant lines would, therefore, be a promising strategy to increase bioavailability of Fe, and also Zn (Petry et al., 2016; Raboy, 2020).

Efficient Fe accumulation in a bioavailable form is an interesting area to explore. Biofortification of edible plants by overexpression of a native ferritin gene applying the GMO technology is an interesting strategy to increase the Fe content in bean seeds. But, as shown in banana, a high ferritin concentration has to be achieved sufficient for food fortification (Yadav et al., 2017). In vacuoles, small Fe and Zn binding molecules, such as nicotianamine and organic acids (malate and citrate), have been further found as pos- sible further targets for mineral biofortification (Hoppler et al., 2014). Mineral (Fe) bioavailability can, however, also be enhanced through improved processing procedures in- cluding soaking, thermal treatments, fermentation and/ or germination. Combining popular traditions with inno- vative treatments, such as germination, is, therefore, an interesting alternative strategy to pursue. Germination and fermentation are thereby useful for increasing the ac- tivity of polyphenol-degrading enzymes and endogenous phytases, which limit the PA content (Carbonaro et al., 2001). Heating promotes, for example, denaturation and hydrolysis of proteins, influencing chelating capacity and significantly modifies the bio-accessibility of minerals (de Oliveira et al., 2018). More research is, however, needed to establish the effect of such processing procedures on mineral (Fe) bioavailability.

Modification in protein solubility and digestibility is a further process which can be explored. Such modification will affect mineral bioavailability (Carbonaro et al., 2005; Iddir et al., 2019). Diet modelling would, likewise, be an

excellent approach to also capture the complexity of a diet as a whole (Mertens et al., 2017). Another important com- ponent is evaluating the effect of traditional cooking prac- tices on the chemical- and nutritional composition of any selected bean lines. Further, evaluating how ingredients of traditional recipes can contribute to the composition of a balanced and a high nutritional-quality dish with a particular emphasis on minerals would be interesting to explore (Durazzo et al., 2019; López et al., 2013).

## 6.3 | Controlling antinutrients

A future challenge is to explore how to obtain more Fe without affecting the amount of antinutrients and of non- essential toxic elements, e.g. cadmium and nickel. These toxic elements, naturally present in trace amounts in the soil, enter the roots via the Fe-regulated transporter-1 me- diated Fe/Zn uptake mechanism (Khan, Bouraine, et al., 2014). If any possible increases in the Fe content will also affect the PA content, particularly under abiotic stress con- ditions, has to be answered. Very few studies have so far investigated this aspect (Campos-Vega et al., 2010; Carbas et al., 2020). In addition, breeding material with specific polyphenol and tannin profiles should be developed with the aim to reduce their negative effect on Fe bioavailabil- ity and to more clearly define their function in Fe bioavail- ability. Also explored should be if antinutritional proteins, such as protease inhibitors, which are expressed as a re- sponse to environmental stress, will influence the Fe con- tent of common bean seeds (Farooq et al., 2018). Giuberti et al. (2019) already found that absence of phaseolin, the main reserve globulin in seeds, with presence of the α- amylase inhibitor is a potential determinant for raising Fe, and also Zn, concentrations in common bean seeds. Introgression of the *lpa* mutation into the above genetic background is thereby an interesting idea to even allow greater improvement of Fe availability.

In summary, any mineral (Fe) optimized beans, which are developed in future bio-fortification programmes, should ultimately also resilient to stresses associated with climatic changes currently threatening future common bean production. Such newly developed common bean varieties should ideally maintain high yields but also have high amounts of minerals while having low amounts of antinutrients, such as PA, under drought/heat stress con- ditions. So far, the impact of drought/heat stress on com- mon bean yield in combination with the effect of stress also on the mineral content of beans has, unfortunately, not been extensively investigated, particularly not under any field conditions. Lack of such field investigations is a major hurdle in the development of common bean va- rieties improved in drought/heat stress tolerance as well
mineral content. Such field investigations are, therefore,
urgently required.

ACKNOWLEDGEMENTS

KK and JV were funded by NRF incentive funding. Partial
funding was also provided to EC from the Knowledge
Hub on Nutrition and Food Security under ERA-NET
ERA-HDHL (No. 696295) for project “SYSTEMIC: An
integrated approach to the challenge of sustainable food
systems: adaptive and mitigatory strategies to address
climate change and malnutrition” and by the European
Regional Development Fund to FS and EC for project
sPATIALS³ under the ROP of the Lombardy Region ERDF
2014–2020—Axis I “Strengthen technological research,
development and innovation”—Action 1.b.1.3 “Support
for co-operative R&D activities to develop new sustainable
technologies, products and services” - Call Hub.

ORCID

Juan Vorster https://orcid.org/0000-0003-3518-3508

Karl Kunert https://orcid.org/0000-0002-7740-3508

REFERENCES

Adhikari, U., Nejadhashemi, A. P., & Woznicki, S. A. (2015). Climate change and eastern Africa: A review of impact on major crops. *Food and Energy Security, 4*, 110–132. https://doi.org/10.1002/fes3.61

Ahmed, S., & Stepp, J. R. (2016). Beyond yields: Climate change effects on specialty crop quality and agroecological management. *Elementa: Science of the Anthropocene, 4*, 92. https://doi.org/10.12952/journal.elementa.000092

Alves Souza, S., Higino Vieira, J., dos Santos, B., Farias, D., da Silva, G. H., & Cabral Aleman, C. (2020). Impact of irrigation frequency and planting density on bean's morpho-physiological and productive traits. *Water, 12*, 2468. https://doi.org/10.3390/w12092468

Andersson, M. S., Saltzman, A., Virk, P. S., & Pfeiffer, W. (2017). Progress update: Crop development of biofortified staple food crops under Harvestplus. *African Journal of Food, Agriculture, Nutrition and Development, 17*, 11906–11935. https://doi.org/10.18697/ajfand.78.HarvestPlus05

Aragão, F. J. L., Barros, L. M. G., De Sousa, M. V., Grossi De Sá, M. F., Almeida, E. R. P., Gander, E. S., & Rech, E. L. (1999). Expression of a methionine-rich storage albumin from the Brazil nut (Bertholletia excelsa HBK Lecythidaceae) in transgenic bean plants (*Phaseolus vulgaris* L. Fabaceae). *Genetics and Molecular Biology, 22*, 445–449. https://doi.org/10.1590/S1415-47571999000300026

Araújo, S. S., Beebe, S., Crespi, M., Delbreil, B., González, E. M., Gruber, V., Lejeune-Henaut, I., Link, W., Monteros, M. J., Prats, E., Rao, I., Vadez, V., & Vaz Patto, M. C. (2015). Abiotic stress responses in legumes: Strategies used to cope with environmental challenges. *Critical Reviews in Plant Sciences, 34*, 237–280. https://doi.org/10.1080/07352689.2014.898450

Asare-Marfo, D., Birol, E., & Gonzalez, C. (2013). Prioritizing countries for biofortification interventions using country-level data.

HarvestPlus Working Paper No. 11. International Food Policy Research Institute (IFPRI).

Assefa, T., Wu, J., Beebe, S. E., Rao, I. M., Marcomin, D., & Claude, R. J. (2015). Improving adaptation to drought stress in small red common bean: Phenotypic differences and predicted genotypic effects on grain yield, yield components and harvest index. *Euphytica, 203*, 477–489. https://doi.org/10.1007/s10681-014-1242-x

Avilés-Gaxiola, S., Chuck-Hernández, C., & Serna Saldívar, S. O. (2018). Inactivation methods of trypsin inhibitor in legumes: A review. *Journal of Food Science, 83*, 17–29. https://doi.org/10.1111/1750-3841.13985

Bailey, R. L., West, K. P. Jr, & Black, R. E. (2015). The epidemiology of global micronutrient deficiencies. *Annals of Nutrition and Metabolism, 66*, 22–33. https://doi.org/10.1159/000371618

Beebe, S. (2012). Common bean breeding in the tropics. *Plant Breeding Reviews, 36*, 357–426. https://doi.org/10.1002/978118358566.ch5

Beebe, S., Gonzalez, A. V., & Rengifo, J. (2000). Research on trace minerals in the common bean. *Food and Nutrition Bulletin, 21*, 387–391. https://doi.org/10.1177/156482650002100408

Beebe, S. E., Ramírez, J., Jarvis, A., Rao, I. M., Mosquera, G., Bueno, J. M., & Blair, M. W. (2011). Genetic improvement of common beans and the challenges of climate change. In S. S. Yadav, R. Redden, J. L. Hatfield, H. Lotze-Campen, & A. Hall (Eds.), *Crop Adaptation to Climate Change* (pp. 356–369). John Wiley & Sons Inc.

Beebe, S. E., Rao, I. M., Blair, M. W., & Acosta-Gallegos, J. A. (2013). Phenotyping common beans for adaptation to drought. *Frontiers in Physiology, 4*, 35. https://doi.org/10.3389/fphys.2013.00035

Beebe, S. E., Rao, I. M., Cajiao, C., & Grajales, M. (2008). Selection for drought resistance in common bean also improves yield in phosphorus limited and favorable environments. *Crop Science, 48*, 582–592. https://doi.org/10.2135/cropsci2007.07.0404

Beebe, S. E., Rao, I. M., Jyostna, D. M., & Polania, J. (2014). Common beans, biodiversity, and multiple stresses: Challenges of drought resistance in tropical soils. *Crop and Pasture Science, 65*, 667–675. https://doi.org/10.1071/CP13303

Bellucci, E., Aguilar, O. M., Alseekh, S., Bett, K., Brezeanu, C., Cook, D., De la Rosa, L., Delledonne, M., Dostatny, D. F., Ferreira, J. J., Geffroy, V., Ghitarrini, S., Kroc, M., Kumar Agrawal, S., Logozzo, G., Marino, M., Mary-Huard, T., McClean, P., Meglič, V., ... Papa, R. (2021). The INCREASE project: Intelligent collections of food-legume genetic resources for European agrofood systems. *Plant Journal, 108*, 646–660. https://doi.org/10.1111/tpj.15472

Bhowmik, P., Konkin, D., Polowick, P., Hodgins, C. L., Subedi, M., Xiang, D., Yu, B., Patterson, N., Rajagopalan, N., Babic, V., Ro, D.-K., Tar'an, B., Bandara, M., Smyth, S. J., Cui, Y., & Kagale, S. (2021). CRISPR/Cas9 gene editing in legume crops: Opportunities and challenges. *Legume Science, 3*(3), 1–16. https://doi.org/10.1002/leg3.96

Bindraban, P. S., Dimkpa, C., Nagarajan, L., Roy, A., & Rabbinge, R. (2015). Revisiting fertilisers and fertilisation strategies for improved nutrient uptake by plants. *Biology and Fertility of Soils, 51*, 897–911. https://doi.org/10.1007/s00374-015-1039-7

Blair, M. W. (2013). Breeding approaches to increasing nutrient-use efficiency. In Z. Rengel (ed.), *Improving Water and Nutrient-Use Efficiency in Food Production Systems*. John
Wiley & Sons, Inc. https://doi.org/10.1002/9781118517994.ch10

Blair, M. W., Fernandez, A. C., Ishitani, M., Moreta, D., Seki, M., Ayling, S., & Shinozaki, K. (2011). Construction and EST sequencing of full-length, drought stress cDNA libraries for common beans (Phaseolus vulgaris L.). *BMC Plant Biology, 11*, 171. https://doi.org/10.1186/1471-2229-11-171

Blair, M. W., Galeano, C. H., Tovar, E., Torres, M. C. M., Castrillón, A. V., Beebe, S. E., & Rao, I. M. (2012). Development of a Mesoamerican intra-genepool genetic map for quantitative trait loci detection in a drought tolerant × susceptible common bean (*Phaseolus vulgaris* L.) cross. *Molecular Breeding, 29*, 71–88. https://doi.org/10.1007/s11032-010-9527-9

Blair, M. W., Herrera, A. L., Sandoval, T. A., Caldas, G. V., Filleppi, M., & Sparvoli, F. (2012). Inheritance of seed phytate and phosphorus levels in common bean (*Phaseolus vulgaris* L.) and association with newly-mapped candidate genes. *Molecular Breeding, 30*, 1265–1277. https://doi.org/10.1007/s11032-012-9713-z

Blair, M. W., Medina, J. I., Astudillo, C., Rengifo, J., Beebe, S. E., Machado, G., & Graham, R. (2010). QTL for seed iron and zinc concentration and content in a Mesoamerican common bean (*Phaseolus vulgaris* L.) population. *Theoretical Applied Genetics, 121*, 1059–1070. https://doi.org/10.1007/s00122-010-1371-0

Boulter, D. (1982). The composition and nutritional value of legumes in relationship to crop improvement by breeding. *Proceedings of the Nutrition Society, 41*, 1–6. https://doi.org/10.1079/PNS19820002

Brear, E. M., Day, D. A., & Smith, P. M. C. (2013). Iron: An essential micronutrient for the legume–rhizobium symbiosis. *Frontiers in Plant Science, 4*, 359. https://doi.org/10.3389/fpls.2013.00359

Broughton, W. J., Hernández, G., Blair, M., Beebe, S., Gepts, P., & Vanderleyden, J. (2003). Beans (Phaseolus spp.) – model food legumes. *Plant and Soil, 252*, 55–128. https://doi.org/10.1023/A:1024146710611

Campion, B., Sparvoli, F., Doria, E., Tagliabue, G., Galasso, I., Fileppi, M., Bollini, R., & Nielsen, E. (2009). Isolation and characterisation of an lpa (low phytic acid) mutant in common bean (*Phaseolus vulgaris* L.). *Theoretical and Applied Genetics, 118*, 1211–1221. https://doi.org/10.1007/s00122-009-0975-8

Campos-Vega, R., Dave Oomah, B., Loarca-Piña, G., & Vergara-Castañeda, H. A. (2013). Common beans and their non-digestible fraction: Cancer inhibitory activity—An Overview. *Foods, 2*, 374–392. https://doi.org/10.3390/foods2030374

Campos-Vega, R., Loarca-Piña, G., & Oomah, B. D. (2010). Minor components of pulses and their potential impact on human health. *Food Research International, 43*, 461–482. https://doi.org/10.1016/j.foodres.2009.09.004

Caproni, L., Raggi, L., Talsma, E. F., Wenzl, P., & Negri, V. (2020). European landrace diversity for common bean biofortification: A genome-wide association study. *Scientific Reports, 10*, 19775. https://doi.org/10.1038/s41598-020-76417-3

Carbas, B., Machado, N., Oppolzer, D., Ferreira, L., Queiroz, M., Brites, C., Rosa, E. A., & Barros, A. I. (2020). Nutrients, antinutrients, phenolic composition, and antioxidant activity of common bean cultivars and their potential for food applications. *Antioxidants (Basel, Switzerland), 9*, 186. https://doi.org/10.3390/antiox9020186

Carbonaro, M. (2021). Nutraceutical perspectives of pulses. In B. K. Tiwari, A. Gowen, & B. Mckenna (Eds.), *Pulse Foods: Processing*, Quality and Nutraceutical Applications (2nd ed., pp. 423–460). Academic Press, Elsevier.

Carbonaro, M., Grant, G., & Cappelloni, M. (2005). Heat-induced denaturation impairs digestibility of legume (*Phaseolus vulgaris* L and *Vicia faba* L) 7S and 11S globulins in the small intestine of rat. *Journal of the Science of Food and Agriculture, 85*, 65–72. https://doi.org/10.1002/jsfa.1940

Carbonaro, M., Grant, G., Mattera, M., Aguzzi, A., & Pusztai, A. (2001). Investigation of the mechanisms affecting Cu and Fe bioavailability from legumes. *Biological Trace Element Research, 84*, 181–196. https://doi.org/10.1385/BTER:84:1-3:181

Carbonaro, M., Marletta, L., & Carnovale, E. (1992). Factors affecting cystine reactivity in proteolytic digests of *Phaseolus vulgaris*. *Journal of Agricultural and Food Chemistry, 40*, 169–174. https://doi.org/10.1021/jf00014a001

Carbonaro, M., Maselli, P., & Nucara, A. (2015). Structural aspects of legume proteins and nutraceutical properties. *Food Research International, 76*, 19–30. https://doi.org/10.1016/j.foodres.2014.11.007

Casañas, F., Pérez-Vega, E., Almirall, A., Plans, M., Sabaté, J., & Ferreira, J. J. (2013). Mapping of QTL associated with seed chemical content in a RIL population of common bean (*Phaseolus vulgaris* L.). *Euphytica, 192*, 279–288. https://doi.org/10.1007/s10681-013-0880-8

Castro-Guerrero, N. A., Isidra-Arellano, M. C., Mendoza-Cozatl, D. G., & Valdés-López, O. (2016). Common bean: A legume model on the rise for unraveling responses and adaptations to iron, zinc, and phosphate deficiencies. *Frontiers in Plant Science, 7*, 600. https://doi.org/10.3389/fpls.2016.00600

Celmeli, T., Sari, H., Canci, H., Sari, D., Adak, A., Eker, T., & Toker, C. (2018). The nutritional content of common bean (*Phaseolus vulgaris* L.) landraces in comparison to modern varieties. *Agronomy, 8*, 166. https://doi.org/10.3390/agronomy8090166

Cerino Badone, F., Amelotti, M., Cassani, E., & Pilu, R. (2012). Study of low phytic acid1-7 (lpa1-7), a new ZmMRP4 mutation in maize. *Journal of Heredity, 103*, 598–605. https://doi.org/10.1093/jhered/ess014

Chekanai, V., Chikowo, R., & Vanlauwe, B. (2018). Response of common bean (*Phaseolus vulgaris* L.) to nitrogen, phosphorus and rhizobia inoculation across variable soils in Zimbabwe. *Agriculture, Ecosystems & Environment, 266*, 167–173. https://doi.org/10.1016/j.agee.2018.08.010

Chiozzotto, R., Ramírez, M., Talbi, C., Cominelli, E., Girard, L., Sparvoli, F., & Hernández, G. (2018). Characterization of the symbiotic nitrogen-fixing common bean low phytic acid (lpa1) mutant response to water stress. *Genes, 9*, 99. https://doi.org/10.3390/genes9020099

Choukri, H., Hejjaoui, K., El-Baouchi, A., El haddad, N., Smouni, A., Maalouf, F., Thavarajah, D., & Kumar, S. (2020). Heat and drought stress impact on phenology, grain yield, and nutritional quality of lentil (*Lens culinaris Medikus*). *Frontiers in Nutrition, 7*, 596307. https://doi.org/10.3389/fnut.2020.596307

Cichy, K. A., Caldas, G. V., Snapp, S. S., & Blair, M. W. (2009). QTL analysis of seed iron, zinc, and phosphorus levels in an Andean bean population. *Crop Science, 49*, 1742–1750. https://doi.org/10.2135/cropsci2008.10.0605

Colombo, F., Paolo, D., Cominelli, E., Sparvoli, F., Nielsen, E., & Pilu, R. (2020). MRP transporters and low phytic acid mutants in major crops: Main pleiotropic effects and future perspectives.
*Frontiers in Plant Science*, **11**, 1301. https://doi.org/10.3389/fpls.2020.01301

Cominelli, E., Confalonieri, M., Carlessi, M., Cortinovis, G., Daminati, M. G., Porch, T. G., Losa, A., & Sparvoli, F. (2018). Phytic acid transport in *Phaseolus vulgaris*: A new low phytic acid mutant in the *PvMRP1* gene and study of the *PvMRPs* promoters in two different plant systems. *Plant Science*, **270**, 1–12. https://doi.org/10.1016/j.plantsci.2018.02.003

Cominelli, E., Galbiati, M., Vavasseur, A., Conti, L., Sala, T., Vuylsteke, M., Leonhardt, N., Dellaporta, S. L., & Tonelli, C. (2005). A guard-cell-specific MYB transcription factor regulates stomatal movements and plant drought tolerance. *Current Biology*, **15**, 1196–1200. https://doi.org/10.1016/j.cub.2005.05.048

Cominelli, E., Galimberti, M., Pongrac, P., Landoni, M., Losa, A., Paolo, D., Daminati, M. G., Bollini, R., Cichy, K. A., Vogel-Mikuš, K., & Sparvoli, F. (2020). Calcium redistribution contributes to the hard-to-cook phenotype and increases PHA-L lectin thermal stability in common bean low phytic acid 1 mutant seeds. *Food Chemistry*, **321**, 126680. https://doi.org/10.1016/j.foodchem.2020.126680

Cominelli, E., Orozco-Arroyo, G., & Sparvoli, F. (2017). Phytic acid biosynthesis and transport in *Phaseolus vulgaris*: Exploitation of new genomic resources. In F. Marsolais, M. Perez de la Vega, & M. Santalla (Eds.), *The Common Bean Genome - Compendium of Plant Genomes* (pp. 167–186). Springer International Publishing. https://doi.org/10.1007/978-3-030-04609-5

Cominelli, E., Pilu, R., & Sparvoli, F. (2020). Phytic acid and transporters: What can we learn from low phytic acid mutants. *Plants*, **9**, 69. https://doi.org/10.3390/plants9010069

Cortés, A. J., Monserrate, F. A., Ramírez-Villegas, J., Madriñán, S., & Blair, M. W. (2013). Drought tolerance in wild plant populations: the case of common beans (*Phaseolus vulgaris* L.). *PLoS One*, **8**, e62898. https://doi.org/10.1371/journal.pone.0062898

Cowling, W. A., Stefanova, K. T., Beeck, C. P., Nelson, M. N., Hargreaves, B. L. W., Sass, O., Gilmour, A. R., & Siddique, K. H. M. (2015). Using the animal model to accelerate response to selection in a self-pollinating crop. *G3: Genes, Genomes, Genetics*, **5**, 1419–1428. https://doi.org/10.1534/g3.115.018838

Curie, C., & Mari, S. (2017). New routes for plant iron mining. *New Phytologist*, **214**, 521–525. https://doi.org/10.1111/nph.14364

Cvitanich, C., Przybyłowicz, W. J., Urbanski, D. F., Jurkiewicz, A. M., Mesjasz-Przybyłowicz, J., Blair, M., Astudillo, C., Jensen, E., & Stougaard, J. (2010). Iron and ferritin accumulate in separate cellular locations in Phaseolus seeds. *BMC Plant Biology*, **10**, 6. https://doi.org/10.1186/1471-2229-10-26

Dahl, W. J., & Stewart, M. L. (2015). Position of the Academy of Nutrition and Dietetics: Health implications of dietary fiber. *Journal of the Academy of Nutrition and Dietetics*, **115**, 1861–1870. https://doi.org/10.1016/j.jand.2015.09.003

Dankher, O. P., & Foyer, C. H. (2018). Climate resilient crops for improving global food security and safety. *Plant Cell & Environment*, **41**, 877–884. https://doi.org/10.1111/pce.13207

Daryanto, S., Wang, L., & Jacinthe, P. A. (2017). Global synthesis of drought effects on cereal, legume, tuber and root crops production: A review. *Agricultural Water Management*, **179**, 18–33. https://doi.org/10.1016/j.agwat.2016.04.022

de Figueiredo, M. A., Boldrin, P. F., Hart, J. J., de Andrade, M. J., Guilherme, L. R., Glahn, R. P., & Li, L. (2017). Zinc and selenium accumulation and their effect on iron bioavailability in

common bean seeds. *Plant Physiology and Biochemistry*, **111**, 193–202. https://doi.org/10.1016/j.plaphy.2016.11.019

De Luque, J. J. R., & Creamer, B. (2014). Major constraints and trends for common bean production and commercialization; establishing priorities for future research. *Agronomia Colombiana*, **32**, 423–431. https://doi.org/10.15446/agron.colomb.v32n3.46052

De Paolis, A., Frugis, G., Giannino, D., Iannelli, M. A., Mele, G., Rugini, E., Silvestri, C., Sparvoli, F., Testone, G., Mauro, M. L., Nicolodi, C., & Caretto, S. (2019). Plant cellular and molecular biotechnology: Following Mariotti's steps. *Plants*, **8**, 18. https://doi.org/10.3390/plants8010018

De Ron, A. M., Rodiño, A. P., Santalla, M., González, A. M., Lema, M. J., Martín, I., & Kigel, J. (2016). Seedling emergence and phenotypic response of common bean germplasm to different temperatures under controlled conditions and in open field. *Frontiers in Plant Science*, **7**, 1087. https://doi.org/10.3389/fpls.2016.01087

DeLaat, D. M., Colombo, C. A., Chiorato, A. F., & Carbonell, S. A. M. (2014). Induction of ferritin synthesis by water deficit and iron excess in common bean (*Phaseolus vulgaris* L.). *Molecular Biology Reports*, **41**, 1427–1435. https://doi.org/10.1007/s11033-013-2987-3

Demelash, B. B. (2018). Common bean improvement status (*Phaseolus vulgaris* L.) in Ethiopia. *Advances in Crop Science and Technology*, **6**, 347. https://doi.org/10.4172/2329-8863.1000347

Dhole, V. J., & Reddy, K. S. (2015). Genetic variation for phytic acid content in mungbean (*Vigna radiata* L. Wilczek). *The Crop Journal*, **3**, 157–162. https://doi.org/10.1016/j.cj.2014.12.002

Diaz, L. M., Ricaurte, J., Tovar, E., Cajiao, C., Terán, H., Grajales, M., Polanía, J., Rao, I., Beebe, S., & Raatz, B. (2018). QTL analyses for tolerance to abiotic stresses in a common bean (*Phaseolus vulgaris* L.) population. *PLoS One*, **13**, e0202342. https://doi.org/10.1371/journal.pone.0202342

Diaz, S., Ariza-Suarez, D., Izquierdo, P., Lobaton, J. D., de la Hoz, J. F., Acevedo, F., Duitama, J., Guerrero, A. F., Cajiao, C., Mayor, V., Beebe, S. E., & Raatz, B. (2020). Genetic mapping for agronomic traits in a MAGIC population of common bean (*Phaseolus vulgaris* L.) under drought conditions. *BMC Genomics*, **21**, 1–20. https://doi.org/10.1186/s12864-020-07213-6

Díaz-Gómez, J., Twyman, R. M., Zhu, C., Farré, G., Serrano, J. C. E., Portero-Otin, M., Muñoz, P., Sandmann, G., Capell, T., & Christou, P. (2017). Biofortification of crops with nutrients: Factors affecting utilization and storage. *Current Opinion in Biotechnology*, **44**, 115–123. https://doi.org/10.1016/j.copbio.2016.12.002

Dramadri, I. O., Nkalubo, S. T., & Kelly, J. D. (2019). Identification of QTL associated with drought tolerance in Andean common bean. *Crop Science*, **59**, 1007–1020. https://doi.org/10.2135/cropsci2018.10.0604

Drewnowski, A. (2010). The nutrient rich foods index helps to identify healthy, affordable foods. *The American Journal of Clinical Nutrition*, **91**, 1095S–1101S. https://doi.org/10.3945/ajcn.2010.28450D

Du, H., Zeng, X., Zhao, M., Cui, X., Wang, Q., Yang, H., Cheng, H., & Yu, D. (2016). Efficient targeted mutagenesis in soybean by TALENs and CRISPR/Cas9. *Journal of Biotechnology*, **217**, 90–97. https://doi.org/10.1016/j.jbiotec.2015.11.005

Durazzo, A., Camilli, E., Marconi, S., Lisciani, S., Gabrielli, P., Gambelli, L., Aguzzi, A., Lucarini, M., Kiefer, J., & Marletta, L. (2019). Nutritional composition and dietary intake of composite dishes traditionally consumed in Italy. *Journal*
of Food Composition and Analysis, 77, 115-124. https://doi.org/10.1016/j.jfca.2019.01.007

Estruch, R., Martínez-González, M. A., Corella, D., Salas-Salvadó, J., Ruiz-Gutiérrez, V., Covas, M. I., Fiol, M., Gómez-Gracia, E., López-Sabater, M. C., Vinyoles, E., Arós, F., Conde, M., Lahoz, C., Lapetra, J., Sáez, G., Ros, E., & PREDIMED Study Investigators (2006). Effects of a Mediterranean-style diet on cardiovascular risk factors: A randomized trial. *Annals of Internal Medicine, 145*, 1-11. https://doi.org/10.7326/0003-4819-145-1-200607040-00004

Fahad, S., Bajwa, A. A., Nazir, U., Anjum, S. A., Farooq, A., Zohaib, A., Sadia, S., Nasim, W., Adkins, S., Saud, S., Ihsan, M. Z., Alharby, H., Wu, C., Wang, D., & Huang, J. (2017). Crop production under drought and heat stress: Plant responses and management options. *Frontiers in Plant Sciences, 8*, 1147. https://doi.org/10.3389/fpls.2017.01147

FAO (2014). *FAOSTAT: Statistics for the year 2014*. Rome. Retrieved from http://faostat3.fao.org/home/E

FAO (2018). *Pulse crops for sustainable farms in sub-Saharan Africa*. Retrieved from http://www.fao.org/3/I8300EN/i8300en.pdf

FAOSTAT (2020). Food and agriculture data: Crops and livestock products. https://www.worldatlas.com/articles/the-world-s-top-dry-bean-producing-countries

Farooq, M., Hussain, M., Usman, M., Farooq, S., Alghamdi, S. S., & Siddique, K. H. M. (2018). Impact of abiotic stresses on grain composition and quality in food legumes. *Journal of Agricultural and Food Chemistry, 66*, 8887-8897. https://doi.org/10.1021/acs.jafc.8b02924

Fenta, B. A., Beebe, S. E., & Kunert, K. J. (2020). Role of fixing nitrogen in common bean growth under water deficit conditions. *Food and Energy Security, 9*, e183. https://doi.org/10.1002/fes3.183

Ferguson, F. (2019). Climate change and abiotic stress mechanisms in plants. *Emerging Topics in Life Sciences, 3*, ETLS20180105.

Fileppi, M., Galasso, I., Tagliabue, G., Daminati, M. G., Campion, B., Doria, E., & Sparvoli, F. (2010). Characterisation of structural genes involved in phytic acid biosynthesis in common bean (*Phaseolus vulgaris* L.). *Molecular Breeding, 25*, 453-470. https://doi.org/10.1007/s11032-009-9344-1

Foyer, C. H., Lam, H.-M., Nguyen, H. T., Siddique, K. H. M., Varshney, R., Colmer, T. D., Cowling, W., Bramley, H., Mori, T. A., Hodgson, J. M., Cooper, J. W., Miller, A. J., Kunert, K., Vorster, J., Cullis, C., Ozga, J. A., Wahlqvist, M. L., Liang, Y., Shou, H., ... Considine, M. J. (2016). Neglecting legumes has compromised global food and nutritional security. *Nature Plants, 2*, 16112. https://doi.org/10.1038/nplants.2016.112

Galbiati, M., Matus, J. T., Francia, P., Rusconi, F., Cañón, P., Medina, C., Conti, L., Cominelli, E., Tonelli, C., & Arce-Johnson, P. (2011). The grapevine guard cell-related VvMYB60 transcription factor is involved in the regulation of stomatal activity and is differentially expressed in response to ABA and osmotic stress. *BMC Plant Biology, 11*, 142. https://doi.org/10.1186/1471-2229-11-142

Ganesan, K., & Xu, B. (2017). Polyphenol-rich dry common beans (*Phaseolus vulgaris* L.) and their health benefits. *International Journal of Molecular Sciences, 18*, 2331. https://doi.org/10.3390/ijms18112331

Gepts, P., Aragão, F., de Barros, E., Blair, M. W., Brondani, R., Broughton, W., Galasso, I., Hernández, G., Kami, J., Lariguet, P., McClean, P., Melotto, M., Miklas, P., Pauls, P., Pedrosa-Harand,

A., Porch, T., Sánchez, F., Sparvoli, F., & Yu, K. (2008). In Genomics of Phaseolus beans, a major source of dietary protein and micronutrients in the tropics. In P. Moore, & R. Ming (Eds.), *Genomics of Tropical Crop Plants* (pp. 113-143). Springer. https://doi.org/10.1007/978-0-387-71219-2_5

Giuberti, G., Tava, A., Mennella, G., Pecetti, L., Masoero, F., Sparvoli, F., Lo Fiego, A., & Campion, B. (2019). Nutrients' and antinutrients' seed content in common bean (*Phaseolus vulgaris* L.) lines carrying mutations affecting seed composition. *Agronomy, 9*, 317. https://doi.org/10.3390/agronomy9060317

Glahn, R., Tako, E., Hart, J., Haas, J., Lung'aho, M., & Beebe, S. (2017). Iron bioavailability studies of the first generation of iron-biofortified beans released in Rwanda. *Nutrients, 9*, 787. https://doi.org/10.3390/nu9070787

Graham, R. D., Welch, R. M., Saunders, D. A., Ortiz-Monasterio, I., Bouis, H. E., Bonierbale, M., de Haan, S., Burgos, G., Thiele, G., Liria, R., Meisner, C. A., Beebe, S. E., Potts, M. J., Kadian, M., Hobbs, P. R., Gupta, R. K., & Twomlow, S. (2007). Nutritious subsistence food systems. *Advances in Agronomy, 92*, 1-74.

Gunjača, J., Carović-Stanko, K., Lazarević, B., Vidak, M., Petek, M., Liber, Z., & Šatović, Z. (2021). Genome-wide association studies of mineral content in common bean. *Frontiers in Plant Sciences, 12*, 636484. https://doi.org/10.3389/fpls.2021.636484

Haas, J. D., Luna, S. V., Lung'aho, M. G., Wenger, M. J., Murray-Kolb, L. E., Beebe, S., Gahutu, J.-B., & Egli, I. M. (2016). Consuming iron biofortified beans increases iron status in Rwandan women after 128 days in a randomized controlled feeding trial. *Journal of Nutrition, 146*, 1586-1592. https://doi.org/10.3945/jn.115.224741

Hageman, A. N., Urban, M. O., & Van Volkenburgh, E. (2020). Sensitivity of leaflet growth rate to drought predicts yield in common bean (*Phaseolus vulgaris*). *Functional Plant Biology, 47*, 792-802. https://doi.org/10.1071/FP19332

Hall, C., Hillen, C., & Garden Robinson, J. (2017). Composition, nutritional value, and health benefits of pulses. *Cereal Chemistry, 94*, 11-31. https://doi.org/10.1094/CCHEM-03-16-0069-FI

Hao, Z., Singh, V. P., & Xia, Y. (2018). Seasonal drought prediction: Advances, challenges, and future prospects. *Reviews of Geophysics, 56*, 108-141. https://doi.org/10.1002/2016R G000549

Hart, J. J., Tako, E., Kochian, L. V., & Glahn, R. P. (2015). Identification of black bean (*Phaseolus vulgaris* L.) polyphenols that inhibit and promote iron uptake by Caco-2 Cells. *Journal of Agriculture and Food Chemistry, 63*, 5950-5956. https://doi.org/10.1021/acs.jafc.5b00531

HarvestPlus (2020). *Getting biofortified food on everyone's plate: HarvestPlus 2019 Annual Report. HarvestPlus Annual Report* 9. International Food Policy Research Institute (IFPRI). http://ebrary.ifpri.org/cdm/ref/collection/p15738coll2/id/133723

Heinemann, A. B., Ramirez-Villegas, J., Souza, T. L. P. O., Didoneta, D., Di Stefano, J. G., Bootee, K. J., & Jarvis, A. (2016). Drought impact on rainfed common bean production areas in Brazil. *Agricultural and Forest Meteorology, 225*, 57-74. https://doi.org/10.1016/j.agrformet.2016.05.010

Heinemann, A. B., Ramirez-Villegas, J., Stone, L. F., & Didonet, A. D. (2017). Climate change determined drought stress profiles in rainfed common bean production systems in Brazil. *Agricultural and Forest Meteorology, 246*, 64-77. https://doi.org/10.1016/j.agrformet.2017.06.005
Herrera, M. D., Acosta-Gallegos, J. A., Reynoso-Camacho, R., & Pérez-Ramírez, I. F. (2019). Common bean seeds from plants subjected to severe drought, restricted-and full-irrigation regimes show differential phytochemical fingerprint. *Food Chemistry*, *294*, 368-377. https://doi.org/10.1016/j.foodc hem.2019.05.076

Hindt, M. N., & Guerinot, M. L. (2012). Getting a sense for signals: Regulation of the plant iron deficiency response. *Biochimica et Biophysica Acta*, *1823*, 1521-1530. https://doi.org/10.1016/j. bbamcr.2012.03.010

Hiza, H. A., Casavale, K. O., Guenther, P. M., & Davis, C. A. (2013). Diet quality of Americans differs by age, sex, race/ethnicity, income, and education level. *Journal of the Academy of Nutrition and Dietetics*, *113*, 297-306. https://doi.org/10.1016/j. jand.2012.08.011

Hoppler, M., Egli, I., Petry, N., Gille, D., Zeder, C., Walczyk, T., Blair, M. W., & Hurrell, R. F. (2014). Iron speciation in beans (*Phaseolus vulgaris*) biofortified by common breeding. *Journal of Food Science*, *79*, C1629-C1634. https://doi.org/10.1111/1750-3841.12548

Hornick, B. A. (2007). Bean consumption by adult women is associated with improved nutrient intakes and healthy eating index scores as demonstrated by menu modeling. *Journal of the Academy of Nutrition and Dietetics*, 107(Supplement 8), A100. https://doi.org/10.1016/j.jada.2007.05.276

Hu, F. B. (2005). Protein, body weight, and cardiovascular health. *The American Journal of Clinical Nutrition*, *82*, 242S-247S. https://doi.org/10.1093/ajcn/82.1.242S

Hummel, M., Hallahan, B. F., Brychkova, G., Ramirez-Villegas, J., Guwela, V., Chataika, B., Curley, E., McKeown, P. C., Morrison, L., Talsma, E. F., Beebe, S., Jarvis, A., Chirwa, R., & Spillane, C. (2018). Reduction in nutritional quality and growing area suitability of common bean under climate change induced drought stress in Africa. *Scientific Reports*, *8*, 1-11. https://doi.org/10.1038/s41598-018-33952-4

Hummel, M., Talsma, E. F., Taleon, V., Londoño, L., Brychkova, G., Gallego, S., Raatz, B., & Spillane, C. (2020). Iron, zinc and phytic acid retention of biofortified, low phytic acid, and conventional bean varieties when preparing common household recipes. *Nutrients*, *12*, 658. https://doi.org/10.3390/nu12030658

Hussain, H. A., Hussain, S., Khaliq, A., Ashraf, U., Anjum, S. A., Men, S., & Wang, L. (2018). Chilling and drought stresses in crop plants: Implications, cross talk, and potential management opportunities. *Frontiers in Plant Science*, *9*, 393. https://doi.org/10.3389/fpls.2018.00393

Iddir, M., Degerli, C., Dingo, G., Desmarchelier, C., Schleeh, T., Borel, P., Larondelle, Y., & Bohn, T. (2019). Whey protein isolate modulates beta-carotene bioaccessibility depending on gastrointestinal digestion conditions. *Food Chemistry*, *291*, 157-166. https://doi.org/10.1016/j.foodchem.2019.04.003

Islam, F. M. A., Basford, K. E., Jara, C., Redden, R. J., & Beebe, S. (2002). Seed compositional and disease resistance differences among gene pools in cultivated common bean. *Genetic Resources and Crop Evolution*, *49*, 285-293. https://doi.org/10.1023/A:1015510428026

Izquierdo, P., Astudillo, C., Blair, M. W., Iqbal, A. M., Raatz, B., & Cichy, K. A. (2018). Meta-QTL analysis of seed iron and zinc concentration and content in common bean (Phaseolus vulgaris L.). *Theoretical Applied Genetics*, *131*, 1645-1658. https://doi.org/10.1007/s00122-018-3104-8

Ji, J., Zhang, C., Sun, Z., Wang, L., Duanmu, D., & Fan, Q. (2019). Genome editing in cowpea Vigna unguiculata using CRISPR-Cas9. *International Journal of Molecular Sciences*, *20*, 2471. https://doi.org/10.3390/ijms20102471

Kaymakanova, M., Stoeva, N., & Mincheva, T. (2008). Salinity and its effects on the physiological response of bean (*Phaseolus vulgaris* L.). *Journal of Central European Agriculture*, *9*, 749-755.

Kazai, P., Noulas, C., Khah, E., & Vlachostergios, D. (2019). Yield and seed quality parameters of common bean cultivars grown under water and heat stress field conditions. *AIMS Agriculture and Food*, *4*, 285-302. https://doi.org/10.3934/agrfood.2019.2.285

Kellogg, W. W., & Schware, R. (2019). *Climate change and society: Consequences of increasing atmospheric carbon dioxide* (p. 192). Routledge. https://doi.org/10.4324/9780429048739

Kelly, J. D., Kolkman, J. M., & Schneider, K. (1998). Breeding for yield in dry bean (*Phaseolus vulgaris* L.). *Euphytica*, *102*, 343-356. https://doi.org/10.1023/A:1018392901978

Kermah, M., Franke, A. C., Adjei-Nsiah, S., Ahiabor, B. D. K., Abaidoo, R. C., & Giller, K. E. (2017). Maize-grain legume intercropping for enhanced resource use efficiency and crop productivity in the Guinea savanna of northern Ghana. *Field Crops Research*, *213*, 38-50. https://doi.org/10.1016/j.fcr.2017.07.008

Khan, G. A., Bouraine, S., Wege, S., Li, Y., de Carbonnel, M., Berthomieu, P., Poirier, Y., & Rouached, H. (2014). Coordination between zinc and phosphate homeostasis involves the transcription factor PHR1, the phosphate exporter PHO1, and its homologue PHO1;H3 in Arabidopsis. *Journal of Experimental Botany*, *65*, 871-884. https://doi.org/10.1093/jxb/ert444

Khan, M. A., Castro-Guerrero, N., & Mendoza-Cozatl, D. G. (2014). Moving toward a precise nutrition: Preferential loading of seeds with essential nutrients over non-essential toxic elements. *Frontiers in Plant Science*, *5*, 51. https://doi.org/10.3389/fpls.2014.00051

Kibido, T., Kunert, K., Makgopa, M., Greve, M., & Vorster, J. (2020). Improvement of rhizobium-soybean symbiosis and nitrogen fixation under drought. *Food and Energy Security*, *9*, e177. https://doi.org/10.1002/fes3.177

Kimani, P. M., & Warsame, A. (2019). Breeding second-generation biofortified bean varieties for Africa. *Food and Energy Security*, *8*, e173. https://doi.org/10.1002/fes3.173

Klein, M., Perfus-Barbeoch, L., Frelet, A., Gaedeke, N., Reinhardt, D., Mueller-Roeber, B., Martinoia, E., & Forestier, C. (2003). The plant multidrug resistance ABC transporter AtMRP5 is involved in guard cell hormonal signalling and water use. *Plant Journal*, *33*, 119-129. https://doi.org/10.1046/j.1365-313X.2003.016012

Kobayashi, T., & Nishizawa, N. K. (2012). Iron uptake, translocation, and regulation in higher plants. *Annual Review of Plant Biology*, *63*, 131-152. https://doi.org/10.1146/annurev-arplant-042811-105522

Kunert, K. J., Vorster, B. J., Fenta, B. A., Kibido, T., Dionisio, G., & Foyer, C. H. (2016). Drought stress responses in soybean roots and nodules. *Frontiers in Plant Science*, *7*, 1015. https://doi.org/10.3389/fpls.2016.01015

Kushi, L. H., Doyle, C., McCullough, M., Rock, C. L., Demark-Wahnefried, W., Bandera, E. V., Gapstur, S., Patel, A. V., Andrews, K., & Gansler, T. (2012). American Cancer Society guidelines on nutrition and physical activity for cancer prevention. *CA: A Cancer Journal for Clinicians*, *62*, 30-67. https://doi.org/10.3322/caac.20140
Kwapata, K., Nguyen, T., & Sticklen, M. (2012). Genetic transformation of common bean (*Phaseolus vulgaris* L.) with the gus color marker, the bar herbicide resistance, and the barley (Hordeum vulgare) HVA1 drought tolerance genes. *International Journal of Agronomy*, 2012, 198960. https://doi.org/10.1155/2012/198960

Lanna, A. C., Silva, R. A., Ferraresi, T. M., Mendonça, J. A., Coelho, G. R. C., Moreira, A. S., Valdisser, P. A. M. R., Brondani, C., & Vianello, R. P. (2018). Physiological characterization of common bean (*Phaseolus vulgaris* L.) under abiotic stresses for breeding purposes. *Environmental Science and Pollution Research*, 25, 31149–31164. https://doi.org/10.1007/s11356-018-3012-0

Latef, A. A. H. A., & Ahmad, P. (2015). Legumes and breeding under abiotic stress. In M. M. Azooz, & P. Ahmad (eds), *Legumes under Environmental Stress* (pp. 60–61). John Wiley & Sons, Ltd. https://doi.org/10.1002/9781118917091.ch1

Lee, J. E., Neumann, M., Duro, D. I., & Schmid, M. (2019). CRISPR-based tools for targeted transcriptional and epigenetic regulation in plants. *PLoS One*, 14, e0222778. https://doi.org/10.1371/journal.pone.0222778

Leisner, C. P. (2020). Review: Climate change impacts on food security- focus on perennial cropping systems and nutritional value. *Plant Science*, 293, 110412. https://doi.org/10.1016/j.plantsci.2020.110412

Leitão, S. T., Bicho, M. C., Pereira, P., Paulo, M. J., Malosetti, M., Araújo, S. S., van Eeuwijk, F., & Vaz Patto, M. C. (2021). Common bean SNP alleles and candidate genes affecting photosynthesis under contrasting water regimes. *Horticulture Research*, 8, 4. https://doi.org/10.1038/s41438-020-00434-6

Li, M. W., Xin, D., Gao, Y., Li, K. P., Fan, K., Muñoz, N. B., Yung, W. S., & Lam, H. M. (2017). Using genomic information to improve soybean adaptability to climate change. *Journal of Experimental Botany*, 68, 1823–1834. https://doi.org/10.1093/jxb/erw348

Lindsey, R. (2020). *Climate change: Atmospheric carbon dioxide*. Retrieved from https://www.climate.gov/news-features/under-standing-climate/climate-change-atmospheric-carbon-dioxide

Lioi, L., & Piergiovanni, A. R. (2013). European common bean genetic and genomic resources of grain legume improvement. In M. Singh, H. D. Upadhyaya, & I. S. Bisht (Eds.), *Genetic and Genomic Resources of Grain Legume Improvement*. Elsevier Inc. https://doi.org/10.1016/B978-0-12-397935-3.00002-5

Liu, J., Gunapati, S., Mihelich, N. T., Stec, A. O., Michno, J. M., & Stupar, R. M. (2019). Genome editing in soybean with CRISPR/Cas9. *Methods in Molecular Biology*, 1917, 217–234. https://doi.org/10.1007/978-1-4939-8991-1_16

Loladze, I. (2014). Hidden shift of the ionome of plants exposed to elevated CO2 depletes minerals at the base of human nutrition, *eLife*, 3, e02245. https://doi.org/10.7554/eLife.02245

Lopes-Caitar, V. S., de Carvalho, M. C. C. G., Darben, L. M., Kuwahara, M. K., Nepomuceno, A. L., Dias, W. P., Abdelnoor, R. V., & Marcelino-Guimarães, F. C. (2013). Genome-wide analysis of the Hsp20 gene family in soybean: Comprehensive sequence, genomic organization and expression profile analysis under abiotic and biotic stresses. *BMC Genomics*, 14, 577. https://doi.org/10.1186/1471-2164-14-577

López, A., El-Naggar, T., Dueñas, M., Ortega, T., Estrella, I., Hernández, T., Gómez-Serranillos, M. P., Palomino, O. M., & Carretero, M. E. (2013). Effect of cooking and germination on phenolic composition and biological properties of dark beans (*Phaseolus vulgaris* L.). *Food Chemistry*, 138, 547–555. https://doi.org/10.1016/j.foodchem.2012.10.107

López-Hernández, F., & Cortés, A. J. (2019). Last-generation genome–environment associations reveal the genetic basis of heat tolerance in common bean (*Phaseolus vulgaris* L.). *Frontiers in Genetics*, 10, 954. https://doi.org/10.3389/fgene.2019.00954

Maillard, A., Diquélou, S., Billard, V., Laîné, P., Garnica, M., Prudent, M., Garcia-Mina, J.-M., Yvin, J.-C., & Ourry, A. (2015). Leaf mineral nutrient remobilization during leaf senescence and modulation by nutrient deficiency. *Frontiers in Plant Science*, 6, 317. https://doi.org/10.3389/fpls.2015.00317

Marcolino-Gomes, J., Rodrigues, F. A., Fuganti-Pagliarini, R., Bendix, C., Nakayama, T. J., Celaya, B., Molinari, H. B. C., Neves de Oliveira, M. C., Harmon, F.-G., & Nepomuceno, A. (2014). Diurnal oscillations of soybean circadian clock and drought responsive genes. *PLoS One*, 9, e86402. https://doi.org/10.1371/journal.pone.0086402

Marentes, E., & Grusak, M. A. (1998). Iron transport and storage within the seed coat and embryo of developing seeds of pea (Pisum sativum L.). *Seed Science Research*, 8, 367–375. https://doi.org/10.1017/S0960258500004293

Mathobo, R., Marais, D., & Steyn, J. M. (2017). The effect of drought stress on yield, leaf gaseous exchange and chlorophyll fluorescence of dry beans (*Phaseolus vulgaris* L.). *Agricultural Water Management*, 180, 118–125. https://doi.org/10.1016/j.agwat.2016.11.005

Matiu, M., Ankerst, D. P., & Menzel, A. (2017). Interactions between temperature and drought in global and regional crop yield variability during 1961–2014. *PLoS One*, 12, e0178339. https://doi.org/10.1371/journal.pone.0178339

Mattei, J., Malik, V., Wedick, N. M., Hu, F. B., Spiegelman, D., Willett, W. C., & Campos, H. (2015). Reducing the global burden of type 2 diabetes by improving the quality of staple foods: The global nutrition and epidemiologic transition initiative. *Global Health*, 11, 23. https://doi.org/10.1186/s12992-015-0109-9

Mbiriri, M., Mukwada, G., & Manatsa, D. (2018). Influence of altitude on the spatiotemporal variations of meteorological droughts in mountain regions of the Free State Province, South Africa. *Advances in Meteorology*, 2018, 5206151. https://doi.org/10.1155/2018/5206151

Medek, D. E., Schwartz, J., & Myers, S. S. (2017). Estimated effects of future atmospheric CO₂ concentrations on protein intake and the risk of protein deficiency by country and region. *Environmental Health Perspectives*, 125, 87002. https://doi.org/10.1289/EHP41

Mendoza-Cózatl, D. G., Xie, Q., Akmakjian, G. Z., Jobe, T. O., Patel, A., Stacey, M. G., Song, L., Demoin, D. W., Jurisson, S. S., Stacey, G., & Scoeder, J. I. (2014). OPT3 is a component of the iron-signaling network between leaves and roots and misregulation of OPT3 leads to an over-accumulation of cadmium in seeds. *Molecular Plant*, 7, 1455–1469. https://doi.org/10.1093/mp/ssu067

Mertens, E., van't Veer, P., Hiddink, G. J., Steijns, J. M. J. M., & Kuijsten, A. (2017). Operationalising the health aspects of sustainable diets: A review. *Public Health Nutrition*, 20, 739–757. https://doi.org/10.1017/S1368980016002664

Mitchell, D. C., Lawrence, F. R., Hartman, T. J., & Curran, J. M. (2009). Consumption of dry beans, peas, and lentils could improve diet quality in the US population. *Journal of the American Dietetic Association*, 109, 909–913. https://doi.org/10.1016/j.jada.2009.02.029

Mittler, R. (2006). Abiotic stress, the field environment and stress combination. *Trends in Plant Science*, 11, 15–19. https://doi.org/10.1016/j.tplants.2005.11.002
Mobley, A. R., Jones, J. M., Rodriguez, J., Slavin, J., & Zelman, K. M. (2014). Identifying practical solutions to meet America's fiber needs: Proceedings from the Food & Fiber Summit. *Nutrients*, **6**, 2540–2551. https://doi.org/10.3390/nu6072540

Mulambu, J., Andersson, M., Palenberg, M., Pfeiffer, W., Saltzman, A., Birol, E., Oparinde, A., Boy, E., Herrington, C., Asare-Marfo, D., Lubobo, A., Mukankusi, C., & Kalubo, S. N. (2017). Iron beans in Rwanda: Crop development and delivery experience. *African Journal of Food, Agriculture, Nutrition and Development*, **17**, 12026–12050. https://doi.org/10.18697/ajfand.HarvestPlus010

Muller, A., Schader, C., El-Hage Scialabba, N., Brüggemann, J., Isensee, A., Erb, K.-H., Smith, P., Klocke, P., Leiber, F., Stolze, M., & Niggli, U. (2017). Strategies for feeding the world more sustainably with organic agriculture. *Nature Communications*, **8**, 1290. https://doi.org/10.1038/s41467-017-01410-w

Müller, B. S. F., Sakamoto, T., Silveira, R. D. D., Zambussi-Carvalho, P. F., Pereira, M., Pappas, G. J. Jr, Costa, M. M., Guimarães, C. M., Pereira, W. J., Brondani, C., & Vianello-Brondani, R. P. (2014). Differentially expressed genes during flowering and grain filling in common bean (Phaseolus vulgaris) grown under drought stress conditions. *Plant Molecular Biology Reporter*, **32**, 438–451. https://doi.org/10.1007/s11105-013-0651-7

Murube, E., Beleggia, R., Pacetti, D., Nartea, A., Frascarelli, G., Lanzavecchia, G., Bellucci, E., Nanni, L., Gioia, T., Marciello, U., Esposito, S., Foresi, G., Logozzo, G., Frega, G. N., Bitocchi, E., & Papa, R. (2021). Characterization of nutritional quality traits of a common bean germplasm collection. *Foods*, **10**, 1572. https://doi.org/10.3390/foods10071572

Myers, S. S., Smith, M. R., Guth, S., Golden, C. D., Vaitla, B., Mueller, N. D., Dangour, A., & Huybers, D. P. (2017). Climate change and global food systems: Potential impacts on food security and undernutrition. *Annual Review of Public Health*, **38**, 259–277. https://doi.org/10.1146/annurev-publhealth-031816-044356

Myers, S. S., Zanobetti, A., Kloog, I., Huybers, P., Leakey, A. D., Bloom, A. J., Carlisle, E., Dietterich, L. H., Fitzgerald, G., Hasegawa, T., Holbrook, N. M., Nelson, R. L., Ottman, M. J., Raboy, V., Sakai, H., Sartor, K. A., Schwartz, J., Seneweera, S., Tausz, M., & Usui, Y. (2014). Increasing CO₂ threatens human nutrition. *Nature*, **510**, 139–142. https://doi.org/10.1038/natur e13179

Nadeem, M., Li, J., Yahya, M., Sher, A., Ma, C., Wang, X., & Qiu, L. (2019). Research progress and perspective on drought stress in legumes: A review. *International Journal of Molecular Sciences*, **20**, 2541. https://doi.org/10.3390/ijms20102541

Nadeem, M. A., Yeken, M. Z., Shahid, M. Q., Habyarimana, E., Yılmaz, H., Alsaleh, A., Hatipoğlu, R., Çilesiz, Y., Khawar, K. M., Ludidi, N., Ercişli, S., Aasim, M., Karaköy, T., & Baloch, F. S. (2021). Common bean as a potential crop for future food security: An overview of past, current and future contributions in genomics, transcriptomics, transgenics and proteomics. *Biotechnology & Biotechnological Equipment*, **35**(1), 758–786. https://doi.org/10.1080/13102818.2021.1920462

Nagy, R., Grob, H., Weder, B., Green, P., Klein, M., Frelet-Barrand, A., Schjoerring, J. K., Brearley, C., & Martinoia, E. (2009). The Arabidopsis ATP-binding cassette protein AtMRP5/AtABCC5 is a high affinity inositol hexakisphosphate transporter involved in guard cell signaling and phytate storage. *Journal of Biological Chemistry*, **284**, 33614–33622. https://doi.org/10.1074/jbc. M109.030247

Nelson, G., Bogard, J., Lividini, K., Arsenault, J., Riley, M., Sulser, T. B., Mason-D'Croz, D., Power, B., Gustafson, D., Herrero, M., Wiebe, K., Cooper, K., Remans, R., & Rosegrant, M. (2018). Income growth and climate change effects on global nutrition security to mid-century. *Nature Sustainability*, **1**, 773–781. https://doi.org/10.1038/s41893-018-0192-z

Oliveira, A. P. D., Mateó, B. D. S. O., Fioroto, A. M., Oliveira, P. V. D., & Naozuka, J. (2018). Effect of cooking on the bioaccessibility of essential elements in different varieties of beans (*Phaseolus vulgaris* L.). *Journal of Food Composition and Analysis*, **67**, 135–140. https://doi.org/10.1016/j.jfca.2018.01.012

Oshone, K., Gebeyehu, S., & Tesfaye, K. (2014). Assessment of common bean (*Phaseolus vulgaris* L.) seed quality produced under different cropping systems by smallholder farmers in Eastern Ethiopia. *African Journal of Food, Agriculture, Nutrition and Development*, **14**, 8566–8584.

Ovando-Martínez, M., Guzmán-Maldonado, S. H., Simsek, S., Bello-Pérez, L. A., & Osorio-Díaz, P. (2014). Effect of water regimes on dietary fiber, polyphenols and antioxidant capacity of black and pinto beans. *Agricultural Sciences*, **5**, 342–352. https://doi.org/10.4236/as.2014.54036

Padhi, E. M. T., & Ramdath, D. D. (2017). A review of the relationship between pulse consumption and reduction of cardiovascular disease risk factors. *Journal of Functional Foods*, **38**, 635–643. https://doi.org/10.1016/j.jff.2017.03.043

Palit, P., Kudapa, H., Zougmore, R., Kholova, J., Whitbread, A., Sharma, M., & Varshney, R. V. (2020). An integrated research framework combining genomics, systems biology, physiology, modelling and breeding for legume improvement in response to elevated CO₂ under climate change scenario. *Current Plant Biology*, **22**, 100149. https://doi.org/10.1016/j.cpb.2020.100149

Panzeri, D., Cassani, E., Doria, E., Tagliabue, G., Forti, L., Campion, B., Bollini, R., Brearley, C. A., Pilu, R., Nielsen, E., & Sparvoli, F. (2011). A defective ABC transporter of the MRP family, responsible for the bean lpa1 mutation, affects the regulation of the phytic acid pathway, reduces seed myo-inositol and alters ABA sensitivity. *New Phytologist*, **191**, 70–83. https://doi.org/10.1111/j.1469-8137.2011.03666.x

Parsons, D. J., Rey, D., Tanguy, M., & Holman, I. P. (2019). Regional variations in the link between drought indices and reported agricultural impacts of drought. *Agricultural Systems*, **173**, 119–129. https://doi.org/10.1016/j.agsy.2019.02.015

Peoples, M. B., Murray John, U., & David, H. (2009). Measuring symbiotic nitrogen fixation by legumes. In D. W. Emerich, & H. B. Krishnan (eds), *Nitrogen fixation in crop production* (pp. 125–170). American Society of Agronomy. DOI: https://doi.org/10.2134/agronmonogr52.c6

Pereira, W. J., Melo, A., Coelho, A., Rodrigues, F. A., Mamidi, S., Alencar, S. A., Lanna, A. C., Valdisser, P. A. M. R., Brondani, C., Nascimento-Júnior, I., Borba, T., & Vianello, R. P. (2020). Genome-wide analysis of the transcriptional response to drought stress in root and leaf of common bean. *Genetics and Molecular Biology*, **43**, e20180259. https://doi.org/10.1590/1678-4685-GMB-2018-0259

Petry, N., Boy, E., Wirth, J. P., & Hurrell, R. F. (2015). Review: The potential of the common bean (*Phaseolus vulgaris*) as a vehicle for iron biofortification. *Nutrients*, **7**, 1144–1173. https://doi.org/10.3390/nu7021144

Petry, N., Egli, I., Gahutu, J. B., Tugirimana, P. L., Boy, E., & Hurrell, R. (2014). Phytic acid concentration influences iron
bioavailability from biofortified beans in Rwandese women with low iron status. *The Journal of Nutrition*, **144**, 1681–1687. https://doi.org/10.3945/jn.115.214254

Petry, N., Egli, I., Zeder, C., Walczyk, T., & Hurrell, R. (2010). Polyphenols and phytic acid contribute to the low iron bioavailability from common beans in young women. *The Journal of Nutrition*, **140**, 1977–1982. https://doi.org/10.3945/jn.110.125369

Petry, N., Rohner, F., Gahutu, J. B., Campion, B., Boy, E., Tugirimana, P. L., Zimmerman, M. B., Zwahlen, C., Wirth, J. P., & Moretti, D. (2016). In Rwandese women with low iron status, iron absorption from low-phytic acid beans and biofortified beans is comparable, but low-phytic acid beans cause adverse gastrointestinal symptoms. *The Journal of Nutrition*, **146**, 970–975. https://doi.org/10.3945/jn.115.223693

Philipo, M., Nbakidemi, P. A., & Mbega, E. R. (2020). Environmental and genotypes influence on seed iron and zinc levels of landraces and improved varieties of common bean (*Phaseolus vulgaris* L.). *Tanzania*, **15**, 100056. https://doi.org/10.1016/j.egg.2020.100056

Polania, J., Poschenrieder, C., Rao, I., & Beebe, S. (2016). Estimation of phenotypic variability in symbiotic nitrogen fixation ability of common bean under drought stress using 15N natural abundance in grain. *European Journal of Agronomy*, **79**, 66–73. https://doi.org/10.1016/j.eja.2016.05.014

Polania, J., Rao, I. M., Cajiao, C., Grajales, M., Rivera, M., Velasquez, F., Raatz, B., & Beebe, S. E. (2017). Shoot and root traits contribute to drought resistance in recombinant inbred lines of MD 23–24 × SEA 5 of common bean. *Frontiers in Plant Science*, **8**, 296. https://doi.org/10.3389/fpls.2017.00296

Polania, J., Rao, I. M., Cajiao, C., Rivera, M., Raatz, B., & Beebe, S. (2016). Physiological traits associated with drought resistance in Andean and Mesoamerican genotypes of common bean (*Phaseolus vulgaris* L.). *Euphytica*, **210**, 17–29. https://doi.org/10.1007/s10681-016-1691-5

Pulecio, J., Verma, N., Mejía-Ramírez, E., Huangfu, D., & Raya, A. (2017). CRISPR/Cas9-based engineering of the epigenome. *Cell Stem Cell*, **21**, 431–447. https://doi.org/10.1016/j stem.2017.09.006

Raboy, V. (2020). Low phytic acid crops: Observations based on four decades of research. *Plants*, **9**, 140. https://doi.org/10.3390/plants9020140

Rainey, K. M., & Griffiths, P. D. (2005). Differential response of common bean genotypes to high temperature. *Journal of the American Society for Horticultural Science*, **130**, 18–23. https://doi.org/10.21273/JASHS.130.1.18

Ramirez-Cabral, N. Y. Z., Kumar, L., & Taylor, S. (2016). Crop niche modeling projects major shifts in common bean growing areas. *Agricultural and Forest Meteorology*, **218**, 102–113. https://doi.org/10.1016/j.agrformet.2015.12.002

Rao, I. M., Miles, J. W., Beebe, S. E., & Horst, W. J. (2016). Root adaptations to soils with low fertility and aluminium toxicity. *Annals of Botany*, **118**, 593–605. https://doi.org/10.1093/aob/mcw073

Rawal, V., & Navarro, D. K. (2019). *The Global of Economy Pulses* (p. 190). Food and Agriculture Organization of the United Nations.

Recchia, G. H., Caldas, D. G. G., Beraldo, A. L. A., Da Silva, M. J., & Tsai, S. M. (2013). Transcriptional analysis of drought-induced genes in the roots of a tolerant genotype of the common bean (*Phaseolus vulgaris* L.). *International Journal of Molecular Sciences*, **14**, 7155–7179. https://doi.org/10.3390/ijms14047155

Redden, R. (2013). New approaches for crop genetic adaptation to the abiotic stresses predicted with climate change. *Agronomy*, **3**, 419–432. https://doi.org/10.3390/agronomy3020419

Rehman, H. M., Cooper, J. W., Lam, H.-M., & Yang, S. H. (2019). Legume biofortification is an underexploited strategy for combatting hidden hunger. *Plant Cell and Environment*, **42**, 52–70. https://doi.org/10.1111/pce.13368

Riquelme, A., Wellmann, E., & Pinto, M. (2007). Effects of ultraviolet-B radiation on common bean (*Phaseolus vulgaris* L.) plants grown under nitrogen deficiency. *Environmental and Experimental Botany*, **60**, 360–367. https://doi.org/10.1016/j.envexbot.2006.12.004

Rosales-Serna, R., Kohashi-Shibata, J., Acosta-Gallegos, J. A., Trejo-López, C., Ortiz-Cereceres, J., & Kelly, J. D. (2004). Biomass distribution, maturity acceleration and yield in drought-stressed common bean cultivars. *Field Crops Research*, **85**, 203–211. https://doi.org/10.1016/S0378-4290(03)00161-8

Roy, S., Liu, W., Sekhar Nandety, R., Crook, A., Mysore, K. S., Pislariu, C. I., Frugoli, J., Dickstein, R., & Udvardia, M. K. (2020). Celebrating 20 years of genetic discoveries in Legume nodulation and symbiotic nitrogen fixation. *The Plant Cell*, **32**, 15–41. https://doi.org/10.1105/tpc.19.00279

Rurangwa, E., Vanlauwe, B., & Giller, K. E. (2018). Benefits of inoculation, P fertilizer and manure on yields of common bean and soybean also increase yield of subsequent maize. *Agriculture, Ecosystems & Environment*, **261**, 219–229. https://doi.org/10.1016/j.agee.2017.08.015

Saboori-Robat, E., Joshi, J., Pajak, A., Solouki, M., Mohsenpour, M., Renaud, J., & Marsolais, F. (2019). Common Bean (*Phaseolus vulgaris* L.) accumulates most S-methylcysteine as its γ-glutamyl dipeptide. *Plants*, **8**, 126. https://doi.org/10.3390/plants8050126

Samago, T. Y., Anniye, E. W., & Dakora, F. D. (2018). Grain yield of common bean (*Phaseolus vulgaris* L.) varieties is markedly increased by rhizobial inoculation and phosphorus application in Ethiopia. *Symbiosis*, **75**, 245–255. https://doi.org/10.1007/s13199-017-0529-9

Samtiya, M., Aluko, R. E., & Dhewa, T. (2020). Plant food anti-nutritional factors and their reduction strategies: An overview. *Food Production, Processing and Nutrition*, **2**, 6. https://doi.org/10.1186/s43014-020-0020-5

Saradadevi, R., Mukankusi, C., Li, L., Amongi, W., Mbiu, J. P., Raatz, B., Ariza, D., Beebe, S., Varshney, R. K., Huttner, E., Kinghorn, B., Banks, R., Rubyogo, J. C., Siddique, K. H. M., & Cowling, W. A. (2021). Multivariate genomic analysis and optimal contributions selection predicts high genetic gains in cooking time, iron, zinc, and grain yield in common beans in East Africa. *Plant Genome*, **14**, e20156. https://doi.org/10.1002/tpg2.20156

Schmutz, J., McClean, P. E., Mamidi, S., Wu, G. A., Cannon, S. B., Grimwood, J., Jenkins, J., Shu, S., Song, Q., Chavarro, C., Torres-Torres, M., Geffroy, V., Moghaddam, S. M., Gao, D., Abernathy, B., Barry, K., Blair, M., Brick, M. A., Chovatia, M., ... Jackson, S. A. (2014). A reference genome for common bean and genome-wide analysis of dual domestications. *Nature Genetics*, **46**, 707–713. https://doi.org/10.1038/ng.3008

Sedlar, A., Kidrič, M., Šuštar-Vozlič, J., Pipan, B., Zadražnik, T., & Meglič, V. (2020). Drought stress response in agricultural plants: A case study of common bean (*Phaseolus vulgaris* L.). In G. Ondrasek (ed.). *Drought - Detection and Solutions*. IntechOpen. DOI: https://doi.org/10.5772/intechopen.86526
Sedlar, A., Zupin, M., Maras, M., Razinger, J., Šuštar-Vozlič, J., Pipan, B., & Meglič, V. (2020). QTL mapping for drought-responsive agronomic traits associated with physiology, phenology, and yield in an Andean intra-gene pool common bean population. *Agronomy*, 10, 225. https://doi.org/10.3390/agronomy10020225

Sehgal, A., Sita, K., Bhandari, K., Kumar, S., Kumar, J., Vara Prasad, P. V., Siddique, K. H. M., & Nayyar, H. (2019). Influence of drought and heat stress, applied independently or in combination during seed development, on qualitative and quantitative aspects of seeds of lentil (Lens culinaris Medikus) genotypes, differing in drought sensitivity. *Plant Cell and Environment*, 42, 198-211. https://doi.org/10.1111/pce.13328

Sehgal, A., Sita, K., Siddique, K. H. M., Kumar, R., Bhogireddy, S., Varshney, R. K., HanumanthaRao, B., Nair, R. M., Prasad, P. V. V., & Nayyar, H. (2018). Drought or/and heat-stress effects on seed filling in food crops: Impacts on functional biochemistry, seed yields, and nutritional quality. *Frontiers in Plant Science*, 9, 1705. https://doi.org/10.3389/fpls.2018.01705

Seidel, S. J., Rachmilevitch, S., Schütze, N., & Lazarovitch, N. (2016). Modelling the impact of drought and heat stress on common bean with two different photosynthesis model approaches. *Environmental Modelling and Software*, 81, 111-121. https://doi.org/10.1016/j.envsoft.2016.04.001

Serra-Majem, L., Roman, B., & Estruch, R. (2006). Scientific evidence of interventions using the Mediterranean diet: A systematic review. *Nutrition Reviews*, 64, S27-S47. https://doi.org/10.1111/j.1753-4887.2006.tb00232.x

Sica, P., Galvao, A., Scariolo, F., Maucieri, C., Nicoletto, C., Pilon, C., Sambo, P., Barcaccia, G., Borin, M., Cabrera, M., & Franklin, D. (2021). Effects of drought on yield and nutraceutical properties of beans (Phaseolus spp.) traditionally cultivated in Veneto, Italy. *Horticulturae*, 7, 17. https://doi.org/10.3390/horticulturae7020017

Sinclair, S. A., & Krämer, U. (2012). The zinc homeostasis network of land plants. *Biochimica et Biophysica Acta*, 1823, 1553-1567. https://doi.org/10.1016/j.bbamcr.2012.05.016

Smith, M. R., Veneklaas, E., Polania, J., Rao, I. M., Beebe, S. E., & Merchant, A. (2019). Field drought conditions impact yield but not nutritional quality of the seed in common bean (*Phaseolus vulgaris* L.). *PLoS One*, 14, e0217099. https://doi.org/10.1371/journal.pone.0217099

Sofi, P. A., Rehman, K., Gull, M., Kumari, J., Djanaguiraman, M., & Prasad, P. V. V. (2021). Integrating root architecture and physiological approaches for improving drought tolerance in common bean (*Phaseolus vulgaris* L.). *Plant Physiology Reports*, 26, 4-221. https://doi.org/10.1007/s40502-021-00570-8

Sørensen, J., & Sessitsch, A. (2007). Plant-associated bacteria Lifestyles and molecular interactions. In J. D. van Elsas, J. K. Jansson, & J. T. Trevors (Eds.), *Modern Soil Microbiology* (pp. 211-236). CRC Press.

Sparvoli, F., & Cominelli, E. (2015). Seed biofortification and phytic acid reduction: A conflict of interest for the plant? *Plants*, 4, 728-755. https://doi.org/10.3390/plants4040728

Steiner, J. L., Briske, D. D., Brown, D. P., & Rottler, C. M. (2018). Vulnerability of *Southern Plains* agriculture to climate change. *Climatic Change*, 146, 201-218. https://doi.org/10.1007/s10584-017-1965-5

Svetleva, D., Velcheva, M., & Bhowmik, G. (2003). Biotechnology as a useful tool in common bean (*Phaseolus vulgaris* L.) improvement. *Euphytica*, 131, 189-200. https://doi.org/10.1023/A:1023983831582

Tako, E., Beebe, S. E., Reed, S., Hart, J. J., & Glahn, R. P. (2014). Polyphenolic compounds appear to limit the nutritional benefit of biofortified higher iron black bean (*Phaseolus vulgaris* L.). *Nutrition Journal*, 13, 28. https://doi.org/10.1186/1475-2891-13-28

Teran, J. C. B. M., Konzen, E. R., Palkovic, A., Tsai, S. M., Rao, I. M., Beebe, S., & Gepts, P. (2019). Effect of drought stress on the genetic architecture of photosynthate allocation and remobilization in pods of common bean (*Phaseolus vulgaris* L.), a key species for food security. *BMC Plant Biology*, 19, 1-15. https://doi.org/10.1186/s12870-019-1774-2

Thomine, S., & Vert, G. (2013). Iron transport in plants: Better be safe than sorry. *Current Opinion in Plant Biology*, 16, 322-327. https://doi.org/10.1016/j.pbi.2013.01.003

Tiwari, M., Trivedi, P. K., & Pandey, A. (2020). Emerging tools and paradigm shift of gene editing in cereals, fruits, and horticultural crops for enhancing nutritional value and food security. *Food and Energy Security*, 10, e258. https://doi.org/10.1002/fes3.258

Trapp, J. J., Urrea, C. A., Cregan, P. B., & Miklas, P. N. (2015). Quantitative trait loci for yield under multiple stress and drought conditions in a dry bean population. *Crop Science*, 55, 1596-1607. https://doi.org/10.2135/cropsci2014.11.0792

Trinidad, T. P., Mallillin, A. C., Loyola, A. S., Sagum, R. S., & Encabo, R. R. (2010). The potential health benefits of legumes as a good source of dietary fibre. *British Journal of Nutrition*, 103, 569-574. https://doi.org/10.1017/S0007114509992157

Tura, H., Edwards, J., Gahlaut, V., Garcia, M., Sznajder, B., Baumann, U., Shahinnia, F., Reynolds, M., Langridge, P., Balyan, H. S., Gupta, P. K., Schnurbusch, T., & Fleury, D. (2020). QTL analysis and fine mapping of a QTL for yield-related traits in wheat grown in dry and hot environments. *Theoretical Applied Genetics*, 133, 239-257. https://doi.org/10.1007/s00122-019-03454-6

Valdisser, P. A. M. R., Müller, B. S. F., de Almeida Filho, J. E., Morais Júnior, O. P., Guimarães, C. M., Borba, T. C. O., de Souza, I. P., Zucchi, M. I., Neves, L. G., Coelho, A. S. G., Brondani, C., & Vianello, R. P. (2020). Genome-wide association studies detect multiple QTLs for productivity in Mesoamerican diversity panel of common bean under drought stress. *Frontiers in Plant Science*, 11, 574674. https://doi.org/10.3389/fpls.2020.574674

Van Horn, L., Carson, J. A. S., Appel, L. J., Burke, L. E., Economos, C., Karmally, W., Lancaster, K., Lichtenstein, A. H., Johnson, R. K., Thomas, R. J., Vos, M., Wylie-Rosett, J., & Kris-Etherton, P. (2016). Recommended dietary pattern to achieve adherence to the American Heart Association/American College of Cardiology (AHA/ACC) Guidelines: A Scientific Statement from the American Heart Association. *Circulation*, 134, e505-e529. https://doi.org/10.1161/CIR.0000000000000462

Varshney, R. K., Ojiewo, C., & Monyo, E. (2019). A decade of tropical legumes projects: Development and adoption of improved varieties, creation of market-demand to benefit smallholder farmers and empowerment of national programmes in sub-Saharan Africa and South Asia. *Plant Breeding*, 138, 379-388. https://doi.org/10.1111/pbr.12744

Vasconcelos, I. M., & Oliveira, J. T. A. (2004). Antinutritional properties of plant lectins. *Toxicon*, 44, 385-403. https://doi.org/10.1016/j.toxicon.2004.05.005

Vasconcelos, M. W., Grusak, M. A., Pinto, E., Gomes, A., Ferreira, H., Balázs, B., Centofanti, T., Ntatsi, G., Savvas, D., Karkanis, A., Williams, M., Vandenberg, A., Toma, L., Shrestha, S., Akaichi, F., Oré Barrios, C., Gruber, S., James, E. K., Maluk, M. K. A., & Iannetta, P. (2020). The Biology of Legumes and Their
Agronomic, Economic, and Social Impact. In M. Hasanuzzaman, S. Araújo, & S. Gill (eds), *The Plant Family Fabaceae*. Springer. DOI: https://doi.org/10.1007/978-981-15-4752-2_1

Venezia, M., & Creasey Krainer, K. M. (2021). Current advancements and limitations of gene editing current advancements and limitations of gene editing in orphan crops. *Frontiers in Plant Science, 12*, 742932. https://doi.org/10.3389/fpls.2021.742932

Villordo-Pineda, E., González-Chavira, M. M., Giraldo-Carbajo, P., Acosta-Gallegos, J. A., & Caballero-Pérez, J. (2015). Identification of novel drought-tolerant-associated SNPs in common bean (*Phaseolus vulgaris*). *Frontiers in Plant Science, 6*, 546. https://doi.org/10.3389/fpls.2015.00546

Vlasova, A., Capella-Gutiérrez, S., Rendón-Anaya, M., Hernández-Oñate, M., Minoche, A. E., Erb, I., Câmara, F., Prieto-Barja, P., Corvelo, A., Sanseverino, W., Westergaard, G., Dohm, J. C., Pappas, G. J., Saburido-Alvarez, S., Kedra, D., Gonzalez, I., Cozzuto, L., Gómez-Garrido, J., Aguilar-Morón, M. A., ... Guigó, R. (2016). Genome and transcriptome analysis of the Mesoamerican common bean and the role of gene duplications in establishing tissue and temporal specialization of genes. *Genome Biology, 17*, 32. https://doi.org/10.1186/s13059-016-0883-6

Vogel, E., Donat, M. G., Alexander, L. V., Meinshausen, M., Ray, D. K., Karoly, D., Meinshausen, N., & Frieler, K. (2019). The effects of climate extremes on global agricultural yields. *Environmental Research Letters, 14*, 54010. https://doi.org/10.1088/1748-9326/ab154b

Vucenik, I. (2019). Anticancer properties of inositol hexaphosphate and inositol: An Overview. *Journal of Nutritional Science and Vitaminology (Tokyo)*, 65, S18-S22. https://doi.org/10.3177/jnsv.65.S18

Wagner, S. C. (2011). Biological nitrogen fixation. *Nature Education Knowledge, 3*, 15.

Wang, Y., Wang, Z., Fu, L., Chen, Y., & Fang, J. (2013). Legume consumption and colorectal adenoma risk: A meta-analysis of observational studies. *PLoS One, 8*, e67335. https://doi.org/10.1371/journal.pone.0067335

White, P.J., & Broadley, M. R. (2009). Biofortification of crops with seven mineral elements often lacking in human diets - Iron, zinc, copper, calcium, magnesium, selenium and iodine. *New Phytologist, 182*, 49–84. https://doi.org/10.1111/j.1469-8137.2008.02738.x

Wilker, J. L. (2021). *Symbiotic nitrogen fixation in heirloom, land-race, and modern participatory-bred and conventionally-bred common bean* (*Phaseolus vulgaris* L.). Doctoral dissertation. University of Guelph. https://hdl.handle.net/10214/24091

Wilker, J., Navabi, A., Rajcan, I., Marsolais, F., Hill, B., Torkamaneh, D., & Pauls, K. P. (2019). Agronomic performance and nitrogen fixation of heirloom and conventional dry bean varieties under low-nitrogen field conditions. *Frontiers in Plant Science, 10*, 7–26. https://doi.org/10.3389/fpls.2019.00952

Wilson, M. S., Bulley, S. J., Pisani, F., Irvine, R. F., & Saiardi, A. (2015). A novel method for the purification of inositol phosphates from biological samples reveals that no phytate is present in human plasma or urine. *Open Biology, 5*, 150014. https://doi.org/10.1098/rsob.150014

Wu, J., Wang, L., & Wang, S. (2016). Comprehensive analysis and discovery of drought-related NAC transcription factors

in common bean. *BMC Plant Biology, 16*, 193. https://doi.org/10.1186/s12870-016-0882-5

Wu, L., Chang, Y., Wang, L., Wang, S., & Wu, J. (2021). The aquaporin gene PvXIP1;2 conferring drought resistance identified by GWAS at seedling stage in common bean. *Theoretical and Applied Genetics*. https://doi.org/10.1007/s00122-021-03978-w

Yadav, K., Patel, P., Srivastava, A. K., & Ganapathi, T. R. (2017). Overexpression of native ferritin gene MusaFer1 enhances iron content and oxidative stress tolerance in transgenic banana plants. *PLoS One, 12*, e0188933. https://doi.org/10.1371/journal.pone.0188933

Zadražnik, T., Hollung, K., Egge-Jacobsen, W., Meglič, V., & Šuštar-Vozlič, J. (2013). Differential proteomic analysis of drought stress response in leaves of common bean (*Phaseolus vulgaris* L.). *Journal of Proteomics, 78*, 254–272. https://doi.org/10.1016/j.jprot.2012.09.021

Zargar, S. M., Mahajan, R., Nazir, M., Nagar, P., Kim, S. T., Rai, V., Masi, A., Ahmad, S. M., Shah, R. A., Ganai, N. A., Agrawal, G. K., & Rakwal, R. (2017). Common bean proteomics: Present status and future strategies. *Journal of Proteomics, 169*, 239–248. https://doi.org/10.1016/j.jprot.2017.03.019

Zengin, F. (2013). Physiological behavior of bean (*Phaseolus vulgaris* L.) seedlings under metal stress. *Biological Research, 46*, 79–85. https://doi.org/10.4067/S0716-97602013000100012

Zhang, Z., Lanza, E., Kris-Etherton, P. M., Colburn, N. H., Bagshaw, D., Rovine, M. J., Ulbrecht, J. S., Bobe, G., Chapkin, R. S., & Hartman, T. J. (2010). A high legume low glycemic index diet improves serum lipid profiles in men. *Lipids, 45*, 765–775. https://doi.org/10.1007/s11745-010-3463-7

Zhou, C., Tan, Y., Goßner, S., Li, Y., Shu, Q., & Engel, K. H. (2018). Stability of the metabolite signature resulting from the OsSULTR3;3 mutation in low phytic acid rice (*Oryza sativa* L.) seeds upon cross-breeding. *Journal of Agricultural Food Chemistry, 66*, 9366–9376. https://doi.org/10.1021/acs.jafc.8b03921

Zhu, B., Sun, Y., Qi, L., Zhong, R., & Miao, X. (2015). Dietary legume consumption reduces risk of colorectal cancer: Evidence from a meta-analysis of cohort studies. *Scientific Reports, 5*, 8797. https://doi.org/10.1038/srep08797

Zielińska-Dawidziak, M. (2015). Plant ferritin-a source of iron to prevent its deficiency. *Nutrients, 7*, 1184–1201. https://doi.org/10.3390/nu7021184

Zolina, O., Simmer, C., Belyaev, K., Gulev, S. K., & Koltermann, P. (2013). Changes in the duration of European wet and dry spells during the last 60 years. *Journal of Climate, 26*, 2022–2047. https://doi.org/10.1175/JCLI-D-11-00498.1

**How to cite this article:** Losa, A., Vorster, J., Cominelli, E., Sparvoli, F., Paolo, D., Sala, T., Ferrari, M., Carbonaro, M., Marconi, S., Camilli, E., Reboul, E., Waswa, B., Ekesa, B., Aragão, F., & Kunert, K. (2022). Drought and heat affect common bean minerals and human diet—What we know and where to go. *Food and Energy Security, 11*: e351. https://doi.org/10.1002/fes3.351