Abiotic Stresses: Alteration of Composition and Grain Quality in Food Legumes
Abiotic Stresses: Alteration of Composition and Grain Quality in Food Legumes
Sarkar, Sumi;Khatun, Marium;Era, Farzana Mustafa;Islam, A. K. M. Mominul;Anwar, Md. Parvez;Danish, Subhan;Datta, Rahul;Islam, A. K. M. Aminul
2021-11-04 00:00:00
agronomy Review Abiotic Stresses: Alteration of Composition and Grain Quality in Food Legumes 1 1 1 2 2 Sumi Sarkar , Marium Khatun , Farzana Mustafa Era , A. K. M. Mominul Islam , Md. Parvez Anwar , 3 , 4 , 1 , Subhan Danish * , Rahul Datta * and A. K. M. Aminul Islam * Department of Genetics and Plant Breeding, Bangabandhu Sheikh Mujibur Rahman Agricultural University, Gazipur 1706, Bangladesh; sumisarkarnupur14@gmail.com (S.S.); mariumkhatun6225@gmail.com (M.K.); farzana@bsmrau.edu.bd (F.M.E.) Department of Agronomy, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh; akmmominulislam@bau.edu.bd (A.K.M.M.I.); parvezanwar@bau.edu.bd (M.P.A.) Department of Soil Science, Faculty of Agricultural Sciences and Technology, Bahauddin Zakariya University, Multan 60800, Pakistan Department of Geology and Pedology, Mendel University, Zemedelska 1, 61300 Brno, Czech Republic * Correspondence: sd96850@gmail.com (S.D.); rahulmedcure@gmail.com (R.D.); aminulgpb@bsmrau.edu.bd (A.K.M.A.I.); Tel.: +880-1715-401519 (A.K.M.A.I.) Abstract: Abiotic stresses varyingly affect the grain composition and quality of food legumes. This paper is aimed at discussing the impact of abiotic stresses on the grain composition and quality of food legumes. As protein is the main grain constituent of food legumes for which it is being consumed by humans as a cheap protein source, abiotic stresses such as heat, cold, drought, salinity and heavy metals alter this grain protein content in different dimensions for different food legumes. Moreover, other valuable constituents such as starch, soluble sugar, oil, fatty acid and fiber content are affected Citation: Sarkar, S.; Khatun, M.; Era, differently by the abiotic stresses. The diverse impact of these abiotic stresses ultimately declines F.M.; Islam, A.K.M.M.; Anwar, M.P.; the grain quality and yield of food legumes. As food legumes play a vital role in the nutritional Danish, S.; Datta, R.; Islam, A.K.M.A. diet of millions of people in the world and are occasionally denoted as the meat of poor people, Abiotic Stresses: Alteration of it is important to recognize that the sustainable production of food legumes, even under various Composition and Grain Quality in environmental stresses, has the potential to ensure protein security for people globally. Therefore, Food Legumes. Agronomy 2021, 11, it has become a necessity to improve the productivity and quality of food legumes under abiotic 2238. https://doi.org/10.3390/ stresses through proper crop management and improved breeding strategies, thus enhancing food agronomy11112238 and economic security to the farmers, particularly in the developing countries of the world. Academic Editor: Ryoichi Araki Keywords: abiotic stress; heat stress; drought; salinity; heavy metals; legume; food quality Received: 2 October 2021 Accepted: 1 November 2021 Published: 4 November 2021 1. Introduction Publisher’s Note: MDPI stays neutral The increasing population along with global climate change are generating a great with regard to jurisdictional claims in influence on the agroecosystem and creating various abiotic stresses that are major threats published maps and institutional affil- for global food security. Therefore, one of the major challenges of this era is to maintain a iations. sustainable yield under these stresses and meet the global food demand with nutritional food. Food legumes are one of the major crops that may be included in the cropping system for attaining the nutritional and protein requirements of this growing population, as the protein gap is likely to increase with the increasing population [1]. Copyright: © 2021 by the authors. Food legumes from the second most important plant family Fabaceae, are agriculturally Licensee MDPI, Basel, Switzerland. important nourishing crops provided as a low-cost and rich source of protein to human This article is an open access article beings. In terms of world production, food legumes rank third after cereals and oilseeds, distributed under the terms and having a strong impact on the agro-ecosystem and human nutrition [1]. Nearly 27% of conditions of the Creative Commons global crop production is occupied with food legumes [2]. Food legumes are consumed Attribution (CC BY) license (https:// mostly for edible proteins and oil; those are considered as the major grain quality compo- creativecommons.org/licenses/by/ nents of food legumes. Food legumes are grown in variable climates and abiotic stresses 4.0/). Agronomy 2021, 11, 2238. https://doi.org/10.3390/agronomy11112238 https://www.mdpi.com/journal/agronomy Agronomy 2021, 11, x FOR PEER REVIEW 2 of 23 Agronomy 2021, 11, 2238 2 of 24 stresses such as temperature, drought, salinity and heavy metals can affect the grain com‐ position and quality of food legumes [3]. Grain starch, protein, oil, fatty acids, amino ac‐ such as temperature, drought, salinity and heavy metals can affect the grain composition ids, sugars, dietary fibers, minerals and vitamin contents are considered as the major com‐ and quality of food legumes [3]. Grain starch, protein, oil, fatty acids, amino acids, sugars, ponents of grain composition that help to determine the quality of food legumes [4]. Abi‐ dietary fibers, minerals and vitamin contents are considered as the major components of otic stresses disturb and distinctly change these grain components and the quality of food grain composition that help to determine the quality of food legumes [4]. Abiotic stresses legumes. Heat stress has a damaging effect on the seed yield and the quality of food leg‐ disturb and distinctly change these grain components and the quality of food legumes. umes as the process of entire seed setting such as the development of a male and female Heat stress has a damaging effect on the seed yield and the quality of food legumes as the gametophyte, fertilization and the development of seed is sensitive to heat stress [5]. Cold process of entire seed setting such as the development of a male and female gametophyte, stress is one of the limiting factors for the early sowing of food legumes, as it disrupts the fertilization and the development of seed is sensitive to heat stress [5]. Cold stress is one membrane stability and whole‐grain contents of food legumes [1]. Food legumes are com‐ of the limiting factors for the early sowing of food legumes, as it disrupts the membrane monly grown in rainfed production systems. As a result, food legumes are more suscep‐ stability and whole-grain contents of food legumes [1]. Food legumes are commonly grown tible to drought and the intensity and frequency of drought have been predicted to in‐ in rainfed production systems. As a result, food legumes are more susceptible to drought crease according to global climate models. Drought affects crop growth and becomes and the intensity and frequency of drought have been predicted to increase according to more devastating during reproduction and grain filling, thus decreasing grain yield [6]. global climate models. Drought affects crop growth and becomes more devastating during The productivity of grain legumes is frequently affected by terminal drought. reproduction and grain filling, thus decreasing grain yield [6]. The productivity of grain Food legumes are highly sensitive to salinity stress, particularly at the seedling and legumes is frequently affected by terminal drought. developmental stages [7]. Salinity stress declines water potential due to abundance in Na Food legumes are highly sensitive to salinity stress, particularly at the seedling and and Cl ions in plant tissues resulting in stomatal closure, photosynthesis decline and in‐ developmental stages [7]. Salinity stress declines water potential due to abundance in Na hibition of growth those ultimately affect the grain composition, yield and quality of food and Cl ions in plant tissues resulting in stomatal closure, photosynthesis decline and legumes [8]. Heavy metal is one of the major constraints in food legume production and inhibition of growth those ultimately affect the grain composition, yield and quality of food the maintenance of grain quality. Heavy metals considerably diminish the grain protein legumes [8]. Heavy metal is one of the major constraints in food legume production and content due to a lowered N uptake and supply to the emerging grains [9]. Ultra‐structural the maintenance of grain quality. Heavy metals considerably diminish the grain protein and anatomical changes in plant cells take place due to the uptake and accumulation of content due to a lowered N uptake and supply to the emerging grains [9]. Ultra-structural and heavy anatomical metals atc hanges higher co inncentrations plant cells take as pla place nt phys due iolog to theicuptake al activand ities accumulation such as nutrition of heavy distrib metals ution, nitro at higher gen fix concentrat ation, enzymatic ions as plant activiphysiological ty, photosynthesis, activities funcsuch tion of as p nutrition ollen and distribution, the nutritional nitr qua ogen lity fixation, of seedsenzymatic are adversactivity ely affect , photosynthesis, ed by heavy meta function l stressof [10 pollen ]. Reseand arch the nutritional quality of seeds are adversely affected by heavy metal stress [10]. Research is is needed on the impact of abiotic stresses on food legume grain composition and quality needed for deve onlothe ping impact progra ofms abiotic to improve stresses on thefood grain legume qualitgrain y as well composition as resist and ance quality to abiot foric developing programs to improve the grain quality as well as resistance to abiotic stresses stresses to ensure the adequate global supply of food legumes as the most significant to ensure the adequate global supply of food legumes as the most significant source of source of vegetable proteins. vegetable proteins. 2. Food Legumes 2. Food Legumes Mainly three types of legumes are used, namely forage legumes, food legumes and Mainly three types of legumes are used, namely forage legumes, food legumes and cover crops, whereas food legumes are mostly used as a rich source of protein [1]. Most cover crops, whereas food legumes are mostly used as a rich source of protein [1]. Most of the legume crops are consumed as food in the mature and dry seed form [11]. Food of the legume crops are consumed as food in the mature and dry seed form [11]. Food legumes inhabit a minimum part of the cultivable land of the world, which is mostly con‐ legumes inhabit a minimum part of the cultivable land of the world, which is mostly quered by major cereal crops (e.g., rice, wheat, maize) [12]. The protein demands of the conquered by major cereal crops (e.g., rice, wheat, maize) [12]. The protein demands of the growing population can be fulfilled by the insertion of food legumes into cropping sys‐ growing population can be fulfilled by the insertion of food legumes into cropping systems. tems. Food legumes play an important and diverse role as a nutritious staple of poor peo‐ Food legumes play an important and diverse role as a nutritious staple of poor people ple around the world as an inexpensive source of protein, complex carbohydrates, vita‐ around the world as an inexpensive source of protein, complex carbohydrates, vitamins mins and fiber [13]. Soybeans, peas, peanuts, lentils, different types of beans and chick‐ and fiber [13]. Soybeans, peas, peanuts, lentils, different types of beans and chickpeas are peas are commonly used food legumes (Table 1). commonly used food legumes (Table 1). Table 1. Different kind of food legumes and their uses in human nutrition. Table 1. Different kind of food legumes and their uses in human nutrition. Sources of Images Sl. Common Sl. Common Scientific Sources of Images Picture Scientific Name Major Use [Accessed on 22 May Picture Major Use No Name No Name Name [Accessed on 22 May 2021] 2021] Mainly used for soybean oil. https://zh-prod-1cc738ca-7d3 https://zh‐prod‐ Mainly used for soybean oil. Additionally Additionally used as food b-4a72-b792-20bd8d8fa069 1cc738ca‐7d3b‐4a72‐ used as food products such as soymilk, products such as soymilk, soy .storage.googleapis.com/s3fs- b792‐20bd8d8fa069.stor‐ 1. Soybean Glycine max soy sauce, some beverages and whipped 1. Soybean Glycine max sauce, some beverages and public/styles/max_650$\ age.googleapis.com/s3fs‐ whipped toppings, toppings, soy‐fortified soy-fortified pastas, breakfast times$650/public/2020- 08/ pub‐ pastas, breakfast cereals and soybeans.jpg?itok=DuPfsOBn cereals and bars [14]. lic/styles/max_650×650/p bars [14]. (accessed on 22 May 2021) Agronomy 2021, 11, 2238 3 of 24 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 3 of 23 Table 1. Cont. ublic/2020‐08/soy‐ ublic/2020‐08/soy‐ ublic/2020‐08/soy‐ ublic/2020‐08/soy‐ beans.jpg?itok ublic/2020‐08/soy =DuP‐‐ ublic/2020‐08/soy‐ ublic/2020‐08/soy‐ beans.jpg?itok=DuP‐ Sl. Common Scientific Sources of Images beans.jpg?itok=DuP‐ beans.jpg?itok=DuP‐ beans.jpg?itok ublic/2020 fsOBn‐08/soy =DuP‐‐ Picture Major Use ublic/2020‐08/soy‐ beans.jpg?itok=DuP‐ beans.jpg?itok=DuP‐ fsOBn No Name Name [Accessed on 22 May 2021] fsOBn fsOBn beans.jpg?itok fsOBn =DuP‐ https://ag‐ beans.jpg?itok=DuP‐ fsOBn fsOBn https://ag‐ https://ag‐ https://ag fsOBn ‐ tfoods.co https://ag .za/wp‐‐con‐ fsOBn https://ag https://ag‐‐ Used as a dry pulse and also as a https://agtfoods.co.za/wp- green tfoods.co.za/wp‐con‐ tfoods.co.za/wp‐con‐ tfoods.co.za/wp‐con‐ 2. Chickpea Cicer arietinum Used as a dry pulse and also as a green tfoods.co https://ag tent/up .za/w‐p‐‐con‐ https://ag‐ Used as a dry pulse and also as a green tfoods.co.za/wp‐con‐ Cicer Used Used as aasdry a dry pulse pulse and and also alas so as a green content/uploads/2018/06/ tfoods.co.za/wp‐con‐ 2. Chickpea Cicer arietinum Used as a dry vegetable pulse and [14, al 15s]o. as a green tent/up‐ 2. Chickpea Cicer arietinum Used as a dry pulse and also as a green tent/up‐ 2. 2. Chic Chickpea kpea Cicer arietinum Used as a dry pulse and also as a green tent/up‐ 2. Chickpea Cicer arietinum vegetable [14,15]. tfoods.co loads/2018/06/ tent/up .za/w‐pDesi ‐con‐‐ tfoods.co.za/wp‐con‐ 2. Chickpea Cicer arietinum arietinum a green vegetable vegetable [14 [,1 154,].15]. Desi-Chickpea_600x600_1.jpg tent/up‐ 2. Chickpea Cicer arietinum vegetable [14,15]. tent/up‐ Used as a dry vegetable pulse and [14, al 15s]o. as a green loads/2018/06/Desi‐ Used as a dry pulse and also as a green vegetable [14,15]. loads/2018/06/Desi‐ vegetable [14,15]. loads/2018/06/Desi‐ 2. Chickpea Cicer arietinum Chickpea_600x600_1.jpg tent/up‐ loads/2018/06/Desi‐ 2. Chickpea Cicer arietinum (accessed on 22tent/up May 2021) ‐ loads/2018/06/Desi‐ loads/2018/06/Desi‐ vegetable [14,15]. Chickpea_600x600_1.jpg vegetable [14,15]. 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Grass pea Lathyrus sativus staple food in many countries of Asia and Africa [21]. 0x@2x.jpg?v=1578338252 Africa [21]. ucts/grass_pea_photo_53 ucts/grass_pea 0x@2x.jpg?v=1578338252 _photo_53 Africa [21]. 0x@2x.jpg?v=1578338252 Africa [21]. 0x@2x.jpg?v=1578338252 0x@2x.jpg?v=1578338252 Agronomy 2021, 11, x FOR PEER REVIEW 4 of 23 https://www.thedailyme al.com/sites/de‐ This bean has anticancer potential. It also fault/files/slideshows/16 12. Navy bean Phaseolus vulgaris helps to lower diabetes risk and greater 70994/2173040/21‐ gut health [14]. navy_beans‐Thinkstock‐ Photos‐494876324.jpg Protects the body from free radical dam‐ http://productkg.com/sit 13. Red bean Vigna umbellata age that helps in controlling blood sugar es/default/files/to‐ levels [16]. matnaya‐fasoltalas_0.jpg https://www.foodsafe‐ Red kidney beans are full of folate (vita‐ Red kidney tynews.com/files/2020/07 14. 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Phaseolus vulgaris ous powerful minerals and enzymes that bean loads/2017/03/cranberry‐ help to lower bad cholesterol [18]. beans‐spread‐onto‐cut‐ ting‐with‐solids.jpg https://www.sut‐ It helps to balance sugar level and re‐ tonsbaytrading.com/wp‐ 18. Adzuki bean Vigna angularis duces the risk of diabetes. Improves the content/up‐ strength of bones [20]. loads/2013/06/adzuki‐ beans.jpg It helps to prevent birth defects as it is in‐ http://storage.goog‐ credibly nutritious and an excellent leapis.com/powop‐as‐ Faba bean Agronomy 2021, 11, 2238 5 of 24 19. Vicia faba source of soluble fiber, protein, manga‐ sets/kew_profiles/KPP‐ (Broad bean) nese, copper folate and many other mi‐ CONT_085134_fullsize.j cronutrients [16]. pg https://ju‐ Table 1. Cont. Helps to prevent chronic disease, diseases diesblog.files.word‐ 20. Lima bean Phaseolus lunatus associated with digestion and stimulates Sl. Common Scientific Sources of Images press.com/2010/10/img_ Picture Major Use blood circulation [14]. No Name Name [Accessed on 22 May 2021] 5993.jpg https://cdn.shopify.com/s/ https://cdn.shopify.com/ Grass pea seeds are used as a Grass pea seeds are used as a common files/1/2333/6781/products/ s/files/1/2333/6781/prod‐ Lathyrus common staple food in many Agronomy 2021, 11, x FOR PEER REVIEW 5 of 23 21. Grass pea Lathyrus sativus staple food in many countries of Asia and 21. Grass pea grass_pea_photo_530x@2x.jpg? Agronomy 2021, 11, x FOR PEER REVIEW 5 of 23 Agronomy 2021, 11, x FOR PEER REVIEW 5 of 23 sativus countries of Asia and Africa ucts/grass_pea_photo_53 Africa [21]. v=1578338252 (accessed on 22 Agronomy 2021, 11, x FOR PEER REVIEW 5 of 23 [21]. 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Sword bean Canavalia gladiata system turn , preven over [ts 16 bone –18]. resorption and in‐ /442d44/1253663464/il_fu (accessed on 22 May 2021) jpg hibits bone turn over [16–18]. llxfull.1253663464_bodq. jpg It is a fiber‐rich bean that helps in remov‐ It is a fiber-rich bean that helps ing toxins and waste products in the gut. https://i.et‐ in removing toxins and waste It is a fiber‐rich bean that helps in remov‐ Helps in preventing constipation and ab‐ systatic.com/15567684/r/i products in the gut. Helps in ing toxins and waste products in the gut. https://i.et‐ 32. Jack bean Canavalia. ensiformis dominal distention. The Vitamin C pre‐ l/fff‐ preventing constipation and https://i.etsystatic.com/1556 Helps in preventing constipation and ab‐ systatic.com/15567684/r/i Canavalia. abdominal sent in this distention. bean helps The in defending 7684/r/il/f the def/2833501891 ffdef/2833501891 /il_794xN 32. Jack bean Canavalia. ensiformis dominal distention. The Vitamin C pre‐ l/fff‐ 32. Jack bean ensiformis Vitamin C present in this bean /il_794xN.2833501891_8key.jpg body against disease‐causing microorgan‐ .2833501891_8key.jpg sent in this bean helps in defending the def/2833501891/il_794xN helps in defending the body (accessed on 22 May 2021) isms such as bacteria and viruses [23]. body against disease‐causing microorgan‐ .2833501891_8key.jpg against disease-causing isms such as bacteria and viruses [23]. microorganisms such as 3. Economic Importance of Food Legumes bacteria and viruses [23]. Food legumes are mainly essential to developing countries, as they offer a source of 3. Economic Importance of Food Legumes protein, trace nutrients and calories to people who are not able to afford more pricy nutri‐ 3. Economic Food legu Importance mes are m of ain Food ly essen Legumes tial to developing countries, as they offer a source of tional sources [24]. These are perfect crops for accomplishing developmental goals such protein, trace nutrients and calories to people who are not able to afford more pricy nutri‐ Food legumes are mainly essential to developing countries, as they offer a source as improving the health and nutrition of humans, reducing poverty and enhancing the tional sources [24]. These are perfect crops for accomplishing developmental goals such of protein, trace nutrients and calories to people who are not able to afford more pricy resilience of the ecosystem [25]. Due to the involvement of food legumes such as pulses in as improving the health and nutrition of humans, reducing poverty and enhancing the nutritional sources [24]. These are perfect crops for accomplishing developmental goals nutritional diversity that helps to eliminate hunger and malnutrition, the Food and Agri‐ resilience of the ecosystem [25]. Due to the involvement of food legumes such as pulses in such as improving the health and nutrition of humans, reducing poverty and enhancing culture Organization (FAO) of the United Nations stated 2016 as the International Year of nutritional diversity that helps to eliminate hunger and malnutrition, the Food and Agri‐ the resilience of the ecosystem [25]. Due to the involvement of food legumes such as pulses Pulses [11]. Food legumes can potentially manage sustainable agriculture through the en‐ culture Organization (FAO) of the United Nations stated 2016 as the International Year of in nutritional diversity that helps to eliminate hunger and malnutrition, the Food and hancement of productivity as well as crop diversity and a reduction in the dependency on Pulses [11]. Food legumes can potentially manage sustainable agriculture through the en‐ Agriculture Organization (FAO) of the United Nations stated 2016 as the International Year external inputs, as food legumes have the capabilities of nitrogen (N) fixation by biological hancement of productivity as well as crop diversity and a reduction in the dependency on of Pulses [11]. Food legumes can potentially manage sustainable agriculture through the means, efficient roles in nutrient and water retention, the ability to increase soil organic mat‐ enhancement external inputs, of pr as oductivity food leguas mes well have as cr the op capabilities diversity and of nitrogen a reduction (N) in fixation the dependency by biologic onal ter (SOM) and aid the recovery of soil health by improving soil properties [26]. As one of means, efficient roles in nutrient and water retention, the ability to increase soil organic mat‐ external inputs, as food legumes have the capabilities of nitrogen (N) fixation by biological the most important food legume producing countries, India has started introducing cool‐ means, ter (SOM) effi cient and aid roles the in recovery nutrient of and soil water health retention by improving , the ability soil properties to increase [26]. soil As or ganic one of season food legumes that fit the rice‐fallow ecology to change the rice‐fallow system into a matter the most (SOM) important and aid food the legume recovery producing of soil health countries, by impr India oving has started soil properties introduci [26 ng ]. cool As‐ rice‐food legume system that will help to not only uplift the socio‐economic condition of one of the most important food legume producing countries, India has started introducing season food legumes that fit the rice‐fallow ecology to change the rice‐fallow system into a smallholder rice farmers, ensuring their food and nutritional security, but also to break the cool-season food legumes that fit the rice-fallow ecology to change the rice-fallow system rice‐food legume system that will help to not only uplift the socio‐economic condition of pests and diseases cycle of rice and improve the soil’s structure and fertility through the into a rice-food legume system that will help to not only uplift the socio-economic condition smallholder rice farmers, ensuring their food and nutritional security, but also to break the augmentation of the overall sustainable productivity of the rice‐fallow system [27]. of smallholder rice farmers, ensuring their food and nutritional security, but also to break pests and diseases cycle of rice and improve the soil’s structure and fertility through the the pests and diseases cycle of rice and improve the soil’s structure and fertility through augmentation of the overall sustainable productivity of the rice‐fallow system [27]. 4. Grain Composition of Food Legumes the augmentation of the overall sustainable productivity of the rice-fallow system [27]. The grain of a food legume is composed of protein, dietary fiber, starch or oil in the 4. Grain Composition of Food Legumes 4. Grain Composition of Food Legumes form of energy, macro and micronutrients, vitamins and several bioactive phytochemicals The grain of a food legume is composed of protein, dietary fiber, starch or oil in the such T h as e an gratio inx oidan f a fots o d[1l9] eg.u The me iprot s com ein po c so edntent of p rof ot efoo in,d d ileg etau rymes fibe (T r, sable tarch 2) o rva oiries l in tfrom he fo r20 m– form of energy, macro and micronutrients, vitamins and several bioactive phytochemicals o40% f ene [28 rgy],. macro and micronutrients, vitamins and several bioactive phytochemicals such as such as antioxidants [19]. The protein content of food legumes (Table 2) varies from 20– antioxidants [19]. The protein content of food legumes (Table 2) varies from 20–40% [28]. 40% [28]. Table 2. Percent protein, carbohydrate and lipid present in different food legumes. Table 2. Percent protein, carbohydrate and lipid present in different food legumes. Food Legumes Protein % Carbohydrate% Lipid% Chickpea 21 62.95 6.04 Food Legumes Protein % Carbohydrate% Lipid% Groundnut 26 16.13 49.24 Chickpea 21 62.95 6.04 Lentil 25 63.35 1.06 Groundnut 26 16.13 49.24 Black gram 25 60 1.64 Lentil 25 63.35 1.06 Mung bean 24 62.62 1.15 Black gram 25 60 1.64 Soybean 40 6.4 21.3 Mung bean 24 62.62 1.15 Pea 25 51.3 1.2 Soybean 40 6.4 21.3 Pigeon pea 22 62.78 1.49 Pea 25 51.3 1.2 Cowpea 24 35.5 0.91 Pigeon pea 22 62.78 1.49 Faba bean 29 44.7 1.4 Cowpea 24 35.5 0.91 Faba bean 29 44.7 1.4 Agronomy 2021, 11, 2238 7 of 24 Table 2. Percent protein, carbohydrate and lipid present in different food legumes. Food Legumes Protein % Carbohydrate% Lipid% Chickpea 21 62.95 6.04 Groundnut 26 16.13 49.24 Lentil 25 63.35 1.06 Black gram 25 60 1.64 Mung bean 24 62.62 1.15 Soybean 40 6.4 21.3 Pea 25 51.3 1.2 Pigeon pea 22 62.78 1.49 Cowpea 24 35.5 0.91 Faba bean 29 44.7 1.4 White lupin 38 0.0 10.0 Adzuki bean 20 62.90 0.53 Navy bean 22 60.75 1.50 Lima bean 21 63.38 0.69 Source: Jukanti1 et al. [15,19], Kamboj and Nanda [17], Amarowicz [22], USDA [29], Ge [30]. It also contains oligosaccharides, phytoestrogens, phyto hemagglutinins (lectins), saponins and phenolic compounds that play metabolic roles in humans who consume these foods frequently [22]. The primary phenolic compounds found in a legume seed and seed coats are phenolic acids, condensed tannins and flavonoids [19]. The phenolic compounds are varyingly distributed in different legume seeds (Table 3) and colored legumes are found with more phenolic compounds than uncolored legumes [19]. The total phenolic content (TPC) provides a wide variability in various food legumes and the antioxidant activity of these legumes is directly related to their TPC [31]. Table 3. Variable phenolic compounds present in food legumes. Legume Phenolic Compounds Quantity (g/g) Found in References Hydroxybenzoics 5.69 Dihydroxybenzoic acid 3.68 p-hydroxybenzoic acid 1.48 Protocatechuic acid 0.36 Protocatechuic aldehyde 0.13 2,3,4-trihydroxybenzoic acid 16.9–29.2 Gallic acid 90.9–136.8 Lentil Seed [18,32,33] Vanillic acid 0.59–3.22 Hydroxycinnamics 3.76 Trans-p-coumaroyl malic acid 10.02 Trans-p-coumaroyl glycolic acid 2.88 Trans-p-coumaric acid 5.74 Sinapic acid 1099–2217 chlorogenic acid 159–213 Trans-p-coumaric acid 37.3 Green Trans-p-coumaric acid derivative 6.4 Seed [18,32] lentil Trans-ferulic acid 10.1 Hydroxybenzoics, 84.92 Salicyclic acid 44.89 Vanillic acid 17.01 P-hydroxybenzoic acid 12.20 Pinto Seed [18,32,33] P-hydroxyphenyl acetic acid 8.42 bean Protocatechuic acid 2.40 Hydroxycinnamic acids 36.31 Trans-ferulic acid 11.80 Agronomy 2021, 11, 2238 8 of 24 Table 3. Cont. Legume Phenolic Compounds Quantity (g/g) Found in References Hydroxybenzoics, 21.66 Vanillic acid 10.71 P-hydroxyphenyl acetic acid 6.92 Cannellini [32,33] Seed P-hydroxybenzoic acid 4.30 bean Hydroxycinnamic acids 23.52 Trans-ferulic acid 8.95 Protocatechuic acid 67.6 Crude [18,20,32] Protocatechuic aldehyde 7.71 Adzuki extract Trans-p-coumaric acid 31.3 bean [18,32,33] Seed Trans-p-coumaroyl malic acid 4.57 Gallic 27 Protocatechuic 18.9 Seed coat Cowpea P-hydroxybenzoic 5.81 [32] Ferulic 26.25 Seed Coumaric acid 1.25 Protocatechuic 217 Cranberry [32,34] Seed coat P-hydroxybenzoic acid 239 beans P-hydroxybenzoic acid 19.2 to 60.5 Chickpea Syringic acid 45.9 Seed [18,32,35] Gentisic acid 8.1 to 26.0 Protocatechuic acid 12.1 to 163.5 Pea Seed [32,35] P-hydroxybenzoic acid 45.4 to 101.7 Benzoic acids 57 Seed soybean Protocatechuic acids 44 [32,33] Ferulic acid 95 10.33 Bean kidney P-hydroxybenzoic [32] 10 Sprout 5. Nutritional and Health Benefits Food legumes are essential for the human diet as an important source of nutrients and amino acids, and it has been suggested by the Finnish National Nutrition Council and the Eatwell Guide in the UK to increase the consumption of vegetable protein predominantly from food legumes rather than the consumption of animal protein [35]. Replacing animal protein with vegetable protein has beneficial and significant positive effects on human health such as reducing cholesterol, useful in the diet of diabetics, controlling hypertension, maintaining a healthy weight, improving the health of the cardiovascular system and preventing some cancers [36,37]. The physiological effects of various food legumes differ significantly based on the variability of phytochemicals present in them, as the intake of these phytochemicals may provide various health benefits and protection against several diseases [16]. Food legumes have a comparatively high vitamins and minerals content (Table 4), mainly potassium, calcium, magnesium, zinc, iron and thiamin (vitamin B1) [23]. It has been suggested by several researchers to decrease animal protein consumption and replace it with proteins derived from plants because a positive correlation was found between a high intake of animal protein and a rise in cardiovascular disease, whereas a negative correlation was found between a high intake of plant protein and a reduction in cardiovascular diseases and overall mortality [38]. Agronomy 2021, 11, 2238 9 of 24 Table 4. Vitamins and minerals constituent of different food legumes. Soybean Chickpea Pea (Per Pigeon Pea Groundnut Lentil Mung Bean Faba Bean Vitamins and Grass Pea (Per Cowpea (Per Lupin (Per (Per 100 g (Per 100 g 100 g (Per 100 g (Per 100 g (Per 100 (Per 100 g (Per 100 g Minerals 100 g Seed) 100 g Seed) 100 g Seed) Seed) Seed) Seed) Seed) Seed) g Seed) Seed) Seed) tocopherol 6.5 mg 2.24 mg 0.11 mg - - - - - - 0.08 mg 1.1 mg tocopherol 23.0 mg 10.68 mg 5.0 mg - - - - - - - 15.3 mg Vitamin B1 1.0 mg 0.477 mg 0.7 mg 0.37–0.54 mg 0.345 mg 0.643 mg 0.64 mg 0.87 mg 0.621 mg 0.55 mg 0.32 mg Vitamin B2 0.46 mg 0.212 mg 0.27 mg 0.18–0.27 mg 0.094 mg 0.187 mg 0.135 mg 0.21 mg 0.233 mg 0.23 mg 0.59 mg Vitamin B3 - 1.541 mg - 1.23–2.02 mg - 2.96 mg 12.06 mg 2.6 mg 2.251 mg - - Vitamin B5 - 1.588 mg - 1.44–2.24 mg 0.703 mg 1.26 mg 1.76 mg 2.14 mg - - - Vitamin B6 1.1 mg 0.55 mg 0.12 mg 0.49–0.66 mg 0.171 mg 0.283 mg 0.348 mg 0.5 mg 0.382 mg 0.37 mg 0.4 mg -Carotene - 40.00 mg - 24.08–41.01 g - - - - 68 g - - Vitamin K - 9.00 mg - - 1.7 g - - 5.0 g - - - Calcium 0.21 g 160 mg 0.05 g 0.97–1.03 g - 130 mg 92 mg 35 mg 132 mg 0.14 g 0.24 g Potassium 1.8 g 875.0 mg 1 g 8.75–9.2 g 475 mg 1392 mg 705 mg 677 mg 1246 mg 1.2 g 1.1 g Magnesium 0.22 g 138 mg 0.12 g 1.14–1.24 g 91 mg 183 mg 168 mg 47 mg 189 mg 0.15 g 0.13 g Phosphorus - 366.0 mg - 4.68–5.13 g 267 mg - 376 mg 281 mg 367 mg - - Iron 8.0 mg 5.0 mg 5.2 mg 1.33–1.53 mg 4.29 mg 5.23 mg 4.58 mg 6.51 mg 6.74 mg 6.7 mg 5.4 mg Copper 1.2 mg 0.847 mg 0.66 mg 6.98–7.95 g 0.458 mg 1.057 mg 1.144 mg 0.75 mg 0.941 g 1.1 mg 0.6 mg Zinc 4.2 mg 4.1 mg 3.2 mg 4.35 mg 2.21 mg 2.76 mg 3.27 mg 3.27 mg 2.68 mg 4.1 mg 5.1 mg Selenium 19 g - 1.6 g - - - - 0.1 g 8.2 g 2 g 4.7 g Source: Jukanti1 et al. [15], Arslan [19,21], Celmeli et al. [14], Mathobo et al. [31], Budhathoki et al. [39–46]. Agronomy 2021, 11, 2238 10 of 24 6. Abiotic Stresses Although food legumes grow in diverse climates, different abiotic stresses such as temperature stress, drought, salinity and heavy metals may hamper the grain quality of food legumes [47]. Food legumes contain essential minerals and nutrients essential for human beings and a deficiency of these elements may lead to malnutrition or other health issues in the human body [48]. These essential elements of food legumes are affected and altered by variable abiotic stresses [49,50]. 6.1. Temperature Stress Food legumes can be alienated into two groups based on different growing seasons, specifically warm- or tropical-season and cool-season food legumes [51]. Common beans, black grams, cowpeas, pigeon peas, mung beans, peanuts and soybeans are mainly grown in hot and humid weather and are known as warm-season food legumes [52]. On the other hand, lentils, peas, chickpeas, grass peas, broad beans and dry beans are known as cool-season food legumes [53]. Food legumes exhibit variable levels of sensitivity to high and low-temperature stresses, which diminishes their performance at different growing stages [54]. Both high and low temperatures may act as abiotic stresses for food legumes if the temperature rises or falls beyond the required temperature level needed for the proper growth and development of the food legumes. 6.1.1. High Temperature Mainly, cool-season food legumes are more sensitive to a high temperature than warm-season food legumes and if the temperature rises above the threshold temperature (Table 5), it turns into severe heat stress at particular growth stages [55]. Agronomy 2021, 11, 2238 11 of 24 Table 5. Effect of heat stress on food legumes at different stages of growth. Threshold Heat Stress Food Legumes Growth Stage Effects References Temp. (Day/Night) Lentil 15–30 Reproductive stage 38/23 Reduced electron flow during photosynthesis [56–59] 30–35 Vegetative development Decreased pollen production, impaired photosystem II Peanut Anthesis 38/22 [56,57] Pod and grain yield Reduced photosynthetic activity; impeded electron donation by OEC (Oxygen-Evolving Pea 15–25 Vegetative growth 30/25 Center) of PS II; reduced oxygen evolution and photochemical energy storage; shutting [57,59,60] of PSI reaction center 15–30 Growth Impaired RuBisCO and sucrose metabolism in leaves; disrupted PSII; damaged structure Chickpea 35/16 [56,59] 25 Reproductive growth and functioning of related enzymes and proteins; decreased stigma receptivity Pigeon pea 18–30 Flowering 45/40 Damaged PSII [56,59] Cowpea 18–28 Flowering 36/27 Tapetal cells degeneration and anther indehiscence [56,57,59] 26 Reproductive 38/30 Abscission of flower, reduced reproductive development; pollen germination, pollen tube 23 Post-anthesis 35 Soybean growth and yield; shrunken pollen; damaged PSII; reduced chlorophyll content and [56–61] 30.2 Pollen germination 35 photosynthesis; decreased Fv/Fm 36.1 Pollen tube growth 38/30 Common bean 20–24 Flowering 32/27 Carbon assimilation limited and NADPH supply reduced; reduced photosynthetic rate [59,62] Efficiency of photosynthesis impaired; reduced sucrose in leaves due to decreased Flowering Mung bean [57,59,63] 28–35 >40/25 sucrose synthesizing enzymes and RuBisCO activity Pod development Broad bean 25–35 Flowering 42 Reduced photosynthesis [57,59,64] Black gram 25–35 Flowering 35 Reduced photosynthesis [65] Cytokinin level reduced in seed leading to diminished seed cell numbers and growth Lupin 20–30 Flowering 38 [57,59] rates of seed, reduced seed growth and development processes Agronomy 2021, 11, 2238 12 of 24 Seed filling is intently associated with the whole-plant senescence process and early senescence takes place by heat stress during the seed filling process that enhances the remobilization of assimilating from the source to sink, thus reducing the seed filling du- ration [66]. The grain development of food legumes is affected by heat stress because the tapetum layer of the grain is disintegrated by heat stress, which decreases the nutrient supply to the microspores and such an impairment leads to anther dehiscence prematurely, impedes carbohydrate synthesis and distribution to the grain and develops fractured em- bryos and poor pods, which ultimately reduces the grain yield [59]. Heat stress significantly decreased the yield of lentils by 70% when it was exposed to a heat wave of 35 C for six days, as lentils are a cool seasoned food legume [66]. The grain composition and quality of food legumes is affected by heat stress in many ways, as heat stress mainly affects the reproductive phases (Figure 1). Figure 1. Effect of heat stress on the reproductive stage of food legumes [64]. Heat stress hampers grain composing elements such as sugar, starch, protein, fatty acids and protein (Table 6). It also alters various components accumulating, primarily, in grain-like starch and proteins by preventing the enzymatic processes required for starch and protein synthesis [67]. The temperature of air and soil increases under heat stress, which adversely affect the grain protein content and quality of food legumes [68]. In most of the food legumes, the grain oil content was found to be increased under heat stress, whereas the protein content was found to be decreased [69,70]. The oil content in the grain was increased under heat stress by 20 and 37% in peanuts and soybeans, respectively [71]. However, in kidney beans, the oil content was found to be declined by 23% under heat stress [72]. The fatty acid composition in the grain of food legumes changes due to heat stress. Heat stress considerably enhanced the oleic acid content, whereas the linoleic acid content was found to be decreased in different food legumes [1]. The N and P content of the soybean grain declined when the temperature rose above 40/30 C [73]. A decrease in total nonstructural carbohydrates was found with increasing temperatures and the ratio of soluble sugars to starch was also found to be decreased in various food legumes, particularly in soybeans [74]. Sucrose and oligosaccharides such as the raffinose content in grains increases with an increasing temperature and monosaccharides such as glucose and fructose decrease with an eminent temperature [75]. Agronomy 2021, 11, 2238 13 of 24 Table 6. Alteration in grain composition of food legumes under heat stress. Temperature Increase % (+) or Grain Food Legumes Decrease % ( ) References Control Heat Stress Composition over Control (Day/Night) (Day/Night) Oleic acid +104% 15/30 C 40/30 C Linolenic acid 48.6 Soybean [1,66] 18/13 C 33/28 C Oil content +37% 18/13 C 33/28 C Sucrose 56% Total sugars 24.5% Starch 53% 20/14 C 32/26 C Protein 19.6% [1,66] Peanut Oil content +20% 20/14 C 26/20 C Oleic acid +24% 25 35/16 Soluble proteins +20% Chickpea [1,66,68,76] 25 >32/20 C Sucrose content 9% Kidney bean 28/18 C 34/24 C Oil content 22.7% [1,66,71] 6.1.2. Low Temperature Low-temperature stress or cold stress can be expressed as a temperature that causes injury or irreversible damage to a crop as it falls under the optimum temperature required for the proper growth and development of the crop. Cold stress not only hampers the vegetative stages of food legumes but also alters reproductive growth and grain compo- sitions (Table 7). During the seed germination of food legumes, cold stress enhances the susceptibility to soil-borne diseases, leading to the poor establishment of crops and even the death of seedlings [54,77]. Table 7. Impact of low temperature on some highly important food legumes. Food Legumes Cold Stress Effects Early vegetative phase damage, impaired microsporogenesis and megasporogenesis, loss of pollen germination, inhibition of pollen tube Soybean 1 C for 4, 6 and 8 h growth, abnormal pod formation and seed filling [54] and alteration in starch, protein, fat and fiber composition [78] Early vegetative phase damage, reduction in embryogenesis and poor seed Pea 3 C quality [54] Early vegetative phase damage, impaired microsporogenesis and <10 C; 10 C for megasporogenesis, pollen viability loss, loss of pollen germination, stigma Chickpea 15–30 min receptivity loss, abnormal pod formation [79] and seed filling [54] and alteration in starch, protein, fat and fiber composition [78] Broad bean 5 C for 24 h Early vegetative phase damage and poor seed quality [54] Food legumes grown in cool seasons are mainly sensitive to cold stress, mostly during the formation of a pod and seed filling [78,80]. Carbohydrate metabolism is impaired by cold stress that may lead to the energy deficiency of different reproductive organs such as style, tapetum and endosperm that ultimately causes the sterility of the gametophyte [81]. In various food legumes, it has been well recognized that phenology and grain filling were damaged by cold stress [82]. The grain filling duration and rate reduce under cold stress as grain filling depends on the source–sink relationship that declines under cold stress. The storage of amino acids, minerals and proteins in the grain of food legumes is inhibited by cold stress. In chickpeas, the sugar concentration in the grain increased, whereas storage amino acids, protein, starch, fat and crude fiber accumulation decreased under cold stress [83]. Agronomy 2021, 11, 2238 14 of 24 6.2. Drought Drought is one of the major constraints that limits food legume production, mainly in the arid and semi-arid tropics and the occurrence of drought during the grain development stages is more critical as it causes a significant yield loss [84]. In food legumes, drought highly affects the composition and quality of the grain (Table 8). Abiotic stress, particularly drought, highly influences the grain protein, fat and carbohydrate contents of food legumes. Although, a mild water scarcity during flowering may prefer an increased grain protein content in some food legumes. However, in maximum food legumes, drought reduces the N, P, Fe and Zn content of the grain that ultimately decreases the total grain protein content [85]. The fatty acid composition of a soybean grain was altered by drought that finally altered the total oil composition, oil stability and oil level in the soybean, especially during grain filling [86]. Table 8. Influences of drought stress on growth stages and grain constituents of food legumes. Drought Stress at Food Legumes Effects References Growth Stages Pod development and Lentil Yield reduction by 70 and 24%, respectively [87] reproductive phase Reproductive phase, Yield loss by 49–54, 27–40 and 49–54%, respectivelyGrain Chickpea anthesis and late protein, sodium, potassium and calcium content reduced by 41, [88,89] ripening 33, 25 and 7%, respectively Oil content of grain reduced by 3% and protein content Reproductive phase, Soybean increased by 5% [90–92] pod set and Seed filling Loss of grain yield by 46–71, 45–50 and 42%, respectively Sucrose and starch content reduced in grain by 29–47 and Reproductive, flowering Common bean 18–20% [93] and Pod filling stage Yield loss by 58–87, 49 and 40%, respectively Reproductive and Grain protein content increased by 8 and 3%, respectively Mung bean [94] vegetative stage Yield reduction by 26% Carbohydrate, fat and protein content increased by 4, 5 and Faba bean Grain filling 3–9%, respectively [83] Grain yield loss by 68% Spotted bean Reproductive stage Protein content of grain increased by 6% [87] Black gram (Mash Flowering and Loss of grain yield by 31–57 and 26%, respectively [95] bean) reproductive Reproductive and pod Cowpea Yield loss by 34–66 and 29%, respectively [92] filling Reproductive phase and Pigeon pea Grain yield loss by 40–55 and 42–57% [83] flowering Reduction in soluble sugar, crude fiber and starch in grain by Lupins 15 days after anthesis [83] 18, 11 and 43%, respectively The oil and oleic acid content in soybeans decrease simultaneously when the grain filling period faces drought [96–98]. The oil content of peanuts is influenced by drought, as drought decreases the digestible carbohydrates such as the sucrose, glucose and fructose concentration affecting the composition of fatty acids in the grain through decreasing the unloading of sugars from the stem to the developing seeds [99,100]. During pod filling, a free amino acid pool increased on cowpea grains but the incorporation of these amino acids into the protein chain was suppressed due to drought, which ultimately reduces the protein-amino acid fraction in the grain [76]. The soluble sugars and starch content decreased in the mature grain of the soybean and the common bean, respectively, under drought [100]. The oil contents of the lupin grain dropped by 50–55% under drought. Drought has a distinct effect on the mineral composition of grains of food legumes. In soybeans, the calcium (Ca), phosphorus (P), copper (Cu), manganese (Mn), molybdenum (Mo) and zinc (Zn) concentrations improved under drought, whereas, the sodium (Na), potassium (K) and calcium (Ca) content reduced but the proline content increased in Agronomy 2021, 11, 2238 15 of 24 chickpeas under drought [1]. Under drought, -tocopherol increased in soybean grains by 2–3 fold, which is helpful for preventing the auto-oxidation of a lipid as the tocopherols found in vegetable oils are well-known antioxidants [6]. During the preliminary stage of seed expansion, the seed sink ability reduces due to drought, which results in a decreasing number of endosperm cells and amyloplasts [76]. Acid invertase is a vital enzyme for the seed development of food legumes and its activity decreases due to drought, thus inhibiting sucrose import. As a result, the scarcity of energy sources and prominent levels of abscisic acid (ABA) lead to a poor grain set under drought [101]. 6.3. Salinity Salt stress is one of the major concerns in arid and semi-arid regions, which comprise about 40% of the land area of the earth. It is a significant constraint for food legume production. Salinity stress interrupts grain composition and the quality of food legumes (Table 9) by affecting hormonal interactions, causing a nutritional imbalance, osmotic effects and ionic toxicity [102,103]. Salt stress disturbs the uptake, accumulation and transport of competitive nutrients in food legumes. The nutritional imbalance in legume plants takes place due to the profusion of the sodium (Na ) and chloride (Cl ) ion concentration at the rhizosphere region because these ions interfere with essential nutrients such as N, P, K, Ca, Zn, boron (B), Mg, Cu and iron (Fe) [104]. Salt stress causes an ionic imbalance + 2+ 2+ + 2+ mainly of K and Ca , creating harmful effects on plants [8]. Ca , K and Mg play a vital role in plant photosynthetic activity, but their concentration decreases under higher + + salt contents due to a competitive uptake of Na and K ion flux, resulting in a deficiency of K and significant yield losses [105]. Salt stress highly affects the oil content and grain protein content because of disturbance in nitrate (NO ) uptake and N metabolism of food legumes [106]. A reduction in stigma receptivity, pollen viability and photo assimilates supply during grain filling takes place due to salt stress that eventually reduces the grain yield of food legumes [107]. In mung beans, the total amount of amino acids, protein, carbohydrates and polysaccharides in the grain decreased with the increasing salt stress and the reduction in carbohydrate and polysaccharide contents headed to a reduced photosynthesis, a nutritional imbalance, ion toxicity and hyperosmotic stress [108,109], whereas N uptake was reduced due to the decline in the total amino acids in the grain of the mung bean under salt stress [109]. The K and P concentrations also declined in the grain of the mung bean with increasing salt stress; however, the concentrations of Na, Ca, Mg and chlorine (Cl) increased [110]. Table 9. Effects of salinity stress on the grain composition and quality of food legumes. Food Legumes Concentration of Salt Impacts Grain protein reduction by 29, 60 and 79%, respectively NaCl 3, 6 and 9 dS m Soybean NaCl 9 dS m Oil content of grain reduced by 77% 7 dS m in loam soil and Yield loss around 46% 6.3 dS m clay soil 3 and 3.8 dS m Loss of grain yield by 50 and 69%, respectively 50 and 100 mM Sodium increased by 200 and 271%, respectively Chickpea 50 and 100 mM Potassium decreased by 79.09 and 72.72%, respectively 2 and 9 dS m Sodium increased by 79.80% and Potassium increased by 0.58% NaCl 40 mM Increase in sodium, 51.03%; potassium, 40.31%; and chloride, 58.41% Lentil (cv. 6796) 3.1 and 2 dS m Grain yield loss found to be 100 and 14%, respectively Reduction in grain protein content of 11 and 20%, respectively Reduction in total soluble sugars of 29 and 32%, respectively Reduction in total amino acids of 19 and 21%, respectively 4500 and 6000 ppm Mung bean Nitrogen content in grain decreased by 37 and 24%, respectively Grain phosphorus content decreased by 30 and 20%, respectively Reduction in grain potassium content by 13 and 8%, respectively 250 mM NaCl 80–100% yield loss Agronomy 2021, 11, 2238 16 of 24 Table 9. Cont. Food Legumes Concentration of Salt Impacts Mungbean (cv. 50 mM NaCl Yield loss by 41% Pusavishal) 6.6 dS m in loam soil Total yield loss around 50% 5.6 dS m in clay soil Yield loss by 52% Total carbohydrates of grain reduced by 9.97 and 33.40%, respectively Faba bean Decrease in grain potassium content of 3.30 and 11.57%, respectively 50 and 100 mM Increase in sodium content of around 12.5 and 62.5%, respectively Magnesium content reduction in grain by 28.57% in both salt concentration Pinto bean (cv. Talash) 8 and 12 dS m Reduction in grain yield by 26 and 41% Source: Farooq et al. [1], Torabian et al. [102], Zhou et al. [105], Ghassemi-Golezani et al. [108], Khan et al. [110], Narula et al. [111]. 6.4. Heavy Metals The accumulation of heavy metals such as mercury (Hg), lead (Pb), cadmium (Cd), chromium (Cr), Cu, Zn, arsenic (As) and nickel (Ni) in the soil is a serious constraint for the crops grown in that soil [112,113]. When these heavy metals are present above the optimum level in the rhizosphere zone, they limit the yield and quality of food legumes (Table 10) as well as cause human health concerns through accumulating in the grains of food legumes [114]. The predominant use of heavy metals leads to a decrease in the yield of food legumes and dangerously affects human health through entering into the food chain [9]. Heavy metal toxicity causes weak plant growth, chlorosis, a yield reduction supplemented by decreased nutrient uptake, plant metabolism disorders and a reduced molecular nitrogen-fixing ability [115]. The uptake of mineral nutrients is altered by heavy metals, which inhibits the opening of the stomata by cooperating with plant water balance, thus disturbing the enzymes of the Calvin cycle, carbohydrate metabolism, photosynthesis and, ultimately, reducing the productivity of food legumes [116]. Cd is a heavy metal highly toxic to plants, humans, animals and causes oxidative stress in plants. Agronomy 2021, 11, 2238 17 of 24 Table 10. Impacts of different level of heavy metals on grain constituents of food legumes. Heavy Metals Food Legumes Level of Metals in Soil or Growth Media Effects References Groundnut - Xerophytic anatomical features and reduction in grain quality [1,9,10] Changes in lipid composition and alteration in the structural component of Common bean 5 g mL [117,118] thylakoid membrane 50 M CdCl Chloroplast damage, reduction in grain filling rate Cadmium (Cd) Pea [119] 2.5 mM Decrease in starch content of seeds Chickpea 23 mg kg Decrease in grain protein by 22% [1,9,120] Green gram 24 mg kg Grain protein reduction by 8% [1,9] Soybean 0.1, 0.5 and 1.0 mM Reduction in grain oil by 23, 28 and 33%, respectively [1,9,97,117] Pigeon pea 56 and 112 mg L Reduced photosynthesis up to 50% [1] Grass pea 25, 50, 100, 200 and 300 ppm Chromosomal abnormalities [1,120] Lead (Pb) Chickpea 195 and 390 mg kg Grain proteins increase by 3 and 6%, respectively [1,9] Soybean - Inhibited growth [98,117] Common bean 500 ppm Reduced seed germination up to 48% [117] Chromium (Cr) Chickpea 67.5 and 135 mg kg Grain protein increased by 3% and decreased by 2%, respectively [1,9,120] Green gram 68 and 136 mg kg Increase in grain protein by 7 and 11%, respectively [1,9] Black gram [1] Mung bean [117,120] Reduced 50% seed germination potential, contamination in the entire food 20 ppm Mercury (Hg) Pea [1] chain Lentil [1,120] Soybean 0.1, 0.5 and 1.0 mM Grain oil reduction by 38, 58 and 68%, respectively [1,97,117] Changes in the ultra-structure of chloroplasts, Pea (50 and 75 M) [1,117,120] swelling of starch grains in the stroma Copper (Cu) Chickpea Reduced grain protein of 9 and 18%, respectively [1,116] 66.9 and 143.8 mg kg Cowpea 5 ppm Adversely affected the germination process Green gram 334.5 and 669 mg kg Grain protein reduced by 4 and 5%, respectively [1] 50, 100, 200 and 400 ppm Reduction in seed germination and seedling growth Chickpea [1,116] 290.1 and 580.2 mg kg Reduced grain protein by 2 and 16%, respectively Nickel (Ni) Cowpea 5 ppm adversely influenced the germination process [1] Pigeon pea 1.0 mM 32% reduction in net photosynthesis, decrease enzyme activity [1,10] Cowpea 5 ppm Adversely influenced the germination process [1] Zinc (Zn) Chickpea 4890 and 9780 mg kg Increased grain protein by 10 and 19%, respectively [1,116,120] Peas 12.5–73.3 mg of sodium arsenate kg Caused interference in mineral nutrient balance [1,114,116] Arsenic (As) Considerable inhibition in seed reserves accumulation such as starch, proteins, Chickpea 5 mg kg [1,114,121] sugars and minerals, reduced the quality of seeds Agronomy 2021, 11, 2238 18 of 24 An accumulation of Cd has potential health risks, and it mostly happens due to the consumption of soybeans grown in contaminated areas as soybeans have more potential in absorbing heavy metals compared to other food legumes [119]. However, the detrimental effects on soybean oil content were found to be greater for Hg than Cd. The grain oil content of soybeans was reduced when exposed to higher Cd and Hg concentrations and the oil content reduction rate was higher with an individual metal rather than a combined effort of metals, which emphasizes the antagonistic effect of heavy metals on the grain oil content [1,120]. Heavy metal changes major and minor fatty acids in food legumes; oleic and linoleic acid decreased significantly in soybeans under heavy metal stress, whereas palmitic, linolenic and stearic acid were markedly increased [120]. The starch content of pea seeds decreased when grown in 2.5 mM Cd [121]. Pb is another heavy metal, and its toxic effects mainly depend on how it reacts with functional groups such as carboxyl, sulfhydryl and amine, which results in a reduction in or loss of enzymatic activity vital for cell function. The total soluble sugars, soluble proteins and starch content of the common bean decreased with the increase in the Cd and Pb concentration when extended with 1 1 different concentrations of Cd and Pb (1.5, 2.0, 2.5, 3.0 g kg for Cd and 2, 4, 6, 8 g kg for Pb) compared to control plants [118]. Pea grains store Fe and Zn, while lentils accumulate low levels of Pb. The grain protein content of maximum food legumes decreased with the increase in Cd, Cr, Ni, Zn, Pb and Cu, except for chickpeas and mung beans. A Zn application in chickpeas decreased the grain protein content [9]. The minerals’ uptake, accumulation and nutritional composition of legume seeds and shoots may be altered by As. The nutrient balance of Zn, Mn and Mg in peas was altered by As when exposed to 12.5 to 73.3 mg of sodium arsenate/kg dry weight of soil [117]. The accumulation of seed reserves such as starch, proteins, sugars and minerals was significantly inhibited in chickpeas when grown in As (5 mg/kg of dry soil) compared to the controls, indicating that As prominently reduced the grain quality of chickpeas [121]. 7. Impacts of Abiotic Stresses on Nodulation and Nitrogen Fixation Abiotic stresses affect the nodulation and nitrogen fixation of legumes. Most impor- tantly, the drought stress because the formation, growth and functioning of nodules are being affected when there is a shortage of water in soil [122]. Under drought stress, different factors interfere with the nitrogenase enzymatic activity such as reducing the stock of ATP, reducing the respiration efficiency, altering the pH gradient across the bacteroid membrane and regulating nitrogenase by substrate or gene expression. A Considerable decrease in nitrogen fixation during soil dehydration has been found in many grain legumes such as chickpeas, peas, cowpeas, faba beans, etc. The stunted growth of a nodule and a partially developed root cortex-embedded organ was found when a nodule was subjected to dry conditions. Nitrogen fixation as well as nodule respiration degrades equivalently to the degree of water insufficiency under drought stress [123]. Nodule oxygen permeability re- duces under drought stress. As a result, nodules face a limited ability to carry out oxidative phosphorylation, although maintaining relatively high photosynthesis [124]. On the other hand, salinity is one of the most limiting factors for leguminous nitrogen fixation. Nodule formation significantly decreases under soil salinity, simultaneously reduc- ing the symbiotic nitrogen fixation. Salt stress reduces root hair formation, thus inhibiting infection threads and, ultimately, degrading the number of nodules. This happens because of the deleterious effect of salt stress on the colonization of the legume root, which restricts the Rhizobia bacterial growth [125]. Heavy metals are another constraint for nitrogen fixation by bacteria in a legume plant. Such metals firstly affect the soil microorganisms. The composition and activities of microbes are being changed dramatically by a high concentration of heavy metals in the soil [126]. The morphology, growth and many activities of multiple groups of microorganisms are found to be altered by the heavy metals such as Ni, Cu, Cd, As and Zn [127]. These metals have been found to enhance lipid peroxidation [128], thus creating oxidative stress for both rhizobia and host legumes. The induction of nodal genes was Agronomy 2021, 11, 2238 19 of 24 found to be inhibited by a high concentration of heavy metals, which cause a loss of the N-fixing ability of rhizobia in association with some leguminous hosts [9]. 8. Conclusions The diverse climatic changes are significantly affecting the agroecosystem. Besides abiotic stresses, pandemic situations created by viruses such as COVID-19 have also hampered the economic and agricultural systems globally. Under such a situation, food legumes are the cheapest source of protein acquisition. The consumption of good quality legumes can be a replacement for animal protein. That is why there is considerable scope for exploring these safe protein sources in the cropping pattern. However, grain legumes’ production, grain composition and quality are hindered by several abiotic stresses, as stated in this review. A collection of stress tolerance diverse germplasms, the development of tolerant variety/varieties through plant breeding or advanced biotechnologies and the introduction of suitable agronomic management packages could be helpful to overcome the abiotic stress effects on legumes for their yield and nutritional quality improvement. This review not only provides an overview on the research that has been conducted, but also to identify the areas in which research on grain legumes is still needed in order to mitigate the abiotic stress effects on legumes. Author Contributions: Conceptualization, S.S. and A.K.M.A.I.; methodology, S.S. and A.K.M.A.I.; validation, S.S. and A.K.M.A.I.; formal analysis, S.S., F.M.E. and A.K.M.A.I.; investigation, S.S., F.M.E. and A.K.M.A.I.; resources, S.S., F.M.E. and A.K.M.A.I.; data curation, S.S., F.M.E. and A.K.M.A.I.; writing—original draft preparation, S.S., M.K., F.M.E. and A.K.M.A.I.; writing—review and editing, A.K.M.A.I., M.P.A., S.D., R.D. and A.K.M.M.I.; visualization, S.S., A.K.M.M.I. and A.K.M.A.I.; super- vision, A.K.M.A.I.; project administration, A.K.M.A.I.; funding acquisition, A.K.M.A.I., R.D. and S.D. All authors have read and agreed to the published version of the manuscript. Funding: This research received no external funding. 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