Biofortification in vegetable crops
Abstract
Micronutrient malnutrition, also known as "hidden hunger," continues to be a major global public health concern, particularly in developing countries. It affects one-third of the world's population and is caused by inadequate consumption of essential vitamins and minerals such iron, zinc, iodine, and provitamin A. Increasing the nutritional content of plants using conventional breeding, transgenic techniques, or agronomy methods is known as biofortification, and it has proven to be an economical and long-lasting way to deal with this issue. While staple crops have received most of the focus, horticulture crops like fruits and vegetables have enormous promise because of their natural abundance of vitamins, minerals, and antioxidants. This review discusses recent developments in biofortification techniques like gene editing, molecular breeding, and nanotechnology. Because of their extensive use and natural nutritional diversity, vegetables are perfect candidates for biofortification. With a focus on iron (Fe), zinc (Zn), selenium (Se), provitamin A, and other micronutrients essential for human health, this review summarizes the state of the art regarding vegetable biofortification techniques, advancements, difficulties, and prospects. The method of adding nutrients to food crops, known as "biofortification," offers a long-term, sustainable way to supply more micronutrients. These days, three popular methods are agronomic, conventional, and genetic engineering. Fertilizers used in agronomic biofortification might temporarily raise micronutrient levels. To create plants with the required nutrients, parent lines with high vitamin or mineral levels can be crossed over a number of generations in traditional plant breeding. By transferring desired traits from one organism to another, genetic engineering techniques are used to create new cultivars and increase their value.
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Introduction
The term "Biofortification" comes from two words: the Greek word "bio" means "life," and the Latin word "fortificare" means "to make strong." In order to address the detrimental economic and health effects of vitamin and mineral shortages in humans, it refers to the nutrient enrichment of crops was reported by Kumar et al. (2025) [20] . Nutrients can be enriched by 1.) Fortification, or adding vitamins and minerals to frequently consumed foods during processing to boost their nutritional worth, is one way to enrich nutrients. 2) Using vitamin and mineral supplements is known as supplementation. 3) According to Gomathi et al. (2019) [16] , Biofortification works better than other techniques for adding nutrients to a crop or plant. When the body lacks the vitamins, minerals, and other nutrients necessary for maintaining healthy tissues and organ function, malnutrition results Cichy et al. (2022) [11] . Around the world, malnutrition exacerbates acute anxiety since it causes serious social and health issues is same as A number of strategies aimed at reducing the global burden of disease include nutrition as a key component. The need for higher productivity was met by the green revolution, but quality suffered as a result. The main cause of micronutrient deficiency among the impoverished today is their inability to pay for dietary supplements is similar to Darshan et al. (2022) [14] . In both industrialized and developing nations, micronutrient deficiencies particularly those of iron, zinc, vitamin A, selenium, and iodine are serious public health issues. Vegetables are a good source of dietary fiber, antioxidants, and important vitamins and minerals, but unless they are biofortified, their nutritional density frequently falls short of what vulnerable populations need on a daily basis. Different from post-harvest fortification, biofortification is the process of increasing nutrient concentrations in edible plant portions by breeding, agronomy, or biotechnology (Awuchi et Vandana et al. 2022) [3,30] . The term "bio-fortification" describes genetically boosting food crops' bioavailable mineral content. The development of biofortified crops also increases the crops' growth efficiency on soils that have a depleted or unavailable mineral composition. For crops like vegetables, berries, and melons that have short juvenile periods to reach fruiting stage, breeding plants with increased phytonutrients is easiest Adeyeye et al. (2019) [2] . However, it is a much longer-term strategy for tree-fruit and nuts, which typically require a juvenile period of many years before fruit-set is possible. Finding plant variations with higher phytonutrient content in germplasm collections or current commercial cultivars is one of the alternative approaches Dobosy et al. (2020) [15] . This can find lines that consumers might already find acceptable, or it can find a possible donor parent with the right phytonutrient background for transfer to a more acceptable plant-type for consumption. [22] . It is estimated that billions of individuals low intake of key vitamins and minerals like iron, zinc, vitamin A, and iodine, causing weakened immune systems, stunted cognitive growth, excess mortality among children and mothers, and lower economic productivity. This kind of malnutrition, which is characterized by a lack of essential vitamins and minerals despite adequate calorie intake, is particularly common among women and children in developing nations (Parida et al., 2023) [32] .
What is Biofortification
The creation of micronutrient-dense staple crops vegetables and grains by the use of current biotechnology, traditional plant breeding techniques, and agronomic methods is known as biofortification. This mechanism increases the concentration of vitamins and nutrients derived from plants in the edible organ as the plant grows and develops O'Hare et al. (2015) [31] . This method of breeding nutrients in food crops offers a long-term, affordable, and sustainable supply of sufficient micronutrients. It involves using the proper breeding strategy and biotechnological techniques to enrich edible parts such as grain, straw, roots, fruits, and tubers with minerals and vitamins by Saltzman et al. (2013) [26] . By enhancing the daily adequacy of micronutrient intake by the individual throughout the life cycle, biofortified staple foods can help minimize "hidden hunger" even if they may not contain as many critical vitamins and micronutrients as industrially fortified meals Cichy et al. (2022) [11] . Agronomic methods, traditional breeding methods, and genetic engineering methods are all used in biofortification.
Additionally, biofortification offers a potential way to spread food and technology to a population that is undernourished and may not have easy access to a variety of diets. 5 Biofortification of Vegetables 107 conventionally fortified foods and supplements Ofori et al. (2022). The various forms of biofortification can greatly raise the amount of vitamins and nutrients in the living product edible portion, and the accumulation of these elements occurs as a result of the plant's regular physiological processes Atulkunda et al. (2021) [4] . The low-cost solution for underdeveloped nations and their citizens who cannot afford high-quality food is to breed plants to increase the Bioavailability of micronutrients in the desired edible food. This strategy will therefore assure improved health by mitigating micronutrient deficiency. The improvement of vitamins and micronutrients in staple foods and vegetables will increase micronutrient consumption, especially among the impoverished, which would ultimately result in a decrease in malnutrition Afify et al. (2023) [1] . Increasing phytonutrients and vitamins in a plant with a lesser life cycle which have short juvenile period to reach the flowering stage like in vegetable can be easily achieved. But this will be difficult for the tree-fruit and nuts where the juvenile period is much longer Das et al. (2017) [12] . The principal objective of biofortification in vegetables should be the production of the nutritionally enriched vegetable crop so that the negative economic and health consequences can be overcome by managing mineral and vitamin deficiencies in humans. Among crops cultivated the fruits and vegetables have a rich source of genetic diversity for micronutrients, and hence they can be easily biofortified through conventional breeding, marker-assisted breeding, or genetic engineering Dwividi et al. (2017). Strategies for biofortification of vegetables with the enhancement of mineral and vitamin, reduction of anti-nutrients to increase the bioavailability of micronutrients.
Concept of Biofortification:
Biofortification refers to the enhancement of nutrient concentration and bioavailability in edible plant parts. Unlike food fortification, biofortification targets the crop itself, ensuring long-term nutritional benefits, especially for rural populations with limited access to diverse diets Ikram et al. (2025) [18] . A dvances in genomics, marker-assisted selection, and gene-editing tools such as CRISPR/Cas9 offer new opportunities to accelerate vegetable biofortification. Integration of agronomic and genetic approaches, along with nutrition education, will enhance the impact of biofortified vegetables Cichy et al.
Importance of Biofortification
In addition to providing naturally fortified foods to populations with limited access to commercially marketed fortified foods, biofortification offers a very affordable, sustainable, and long-term way to supply more micronutrients in relatively remote rural locations (Roriz et al., 2020) [25] . [10] . They exhibit significant catalytic activity and a big surface area in relation to their size. Essential plant nutrients including iron, zinc, and manganese are known as nano micronutrients. These nutrients are encapsulated or synthesized into nanoparticles to enhance plant absorption and delivery (AOAC 2026) [6] . Additionally, the term "nano-PGPR" refers to Plant Growth-Promoting Rhizobacteria (PGPR) that have been nanoencapsulated or nano-formulated. This technique employs nanotechnology to increase the efficiency of helpful soil bacteria in agriculture Sharma et al. (2020) [29] .
Methods of Biofortification
Biofortification can be achieved through three strategies Agronomic Biofortification Conventional plant breeding Genetic engineering
Agronomic Biofortification
Application of fertilizers to boost the amount of micronutrients in edible portions. The best micronutrients for agronomic biofortification are selenium (as selenate), zinc (foliar treatments of ZnSO4), and iodine (soil application of iodide or iodate) Roriz et al. (2020) [25] . The quick and simple way to fortify micronutrients (Fe, Zn, Cu, etc.) in plants is through foliar spray. Mycorrhizal relationships raise the concentrations of Fe, Se, Zn, and Cu in crop plants, according to several studies. Micronutrients such as Zn, Cu, and Fe are more efficiently absorbed when AM-fungi are present. Onion sulphur concentration is increased by sulphur-oxidizing bacteria Dobosy et al. (2020) [15] .
Biofortification of crops with Iron
The production of hemoglobin and myoglobin, which are involved in the movement of oxygen in blood and muscles, depends on iron. Anemia, frailty, stunted child growth, and increased maternal mortality are all consequences of iron deficiency Jangra et al. (2025) [19] . High-iron beans (Phaseolus vulgaris), specifically cultivated through traditional breeding methods in East Africa, are a hallmark of biofortified crops for iron (Beebe, 2020) [9] . Due to its inherent high iron content, spinach and amaranth have also been selected to have their iron content increased through variations. Iron-rich leafy greens that can be further improved are pumpkin and cowpea leaves. He also stated that Spirulina platensis has been employed as a biofortifying agent to improve the iron status in Amaranthus gangeticus plants, and that utilizing S. platensis as a microbial inoculant increased the iron levels of Amaranthus plants when compared with control Smolen et al. (2019) [32] .
Biofortification of crops with Zinc
At a foliar Zn treatment rate of 1.08 g plant-1, the association between tuber Zn content and foliar Zn application followed a saturation curve, peaking at around 30 mg Zn kg-1 DM Gomathi et al. (2017) [16] . Even after foliar Zn fertilization with 2.16 g Zn plant-1, the content of Zn in the shoots increased by 40 times compared to the unfertilized controls. The application of "Riverm" fertilizer aids in the zinc enrichment of tomatoes, eggplant, and sweet peppers. Vegetables that have been biofortified had 6.60-8.59% more zinc than control Rahim et al. (2020) [24] .
Biofortification of crops with Selenium
When used as a soil amendment, Se-enriched S. pinnata provides beneficial forms of organic-Se to broccoli and carrots Kalia et al. (2017) [21] . A 77Se (IV) solution that was enriched to 99.7% as 77Se was applied topically to onions and carrots to biofortify them. Selenium treatment had no effect on the production or oil content of brassica crops Adeyeye et al. (2019) [2] . There found a high concentration of Se (1.92-1.96 μg Se g-1) in the meal and seeds.
Conventional plant breeding
The amount of nutrient status in the current varieties has dropped due to traditional breeding's emphasis on yield qualities and resistance breeding during the past 40 years. Fe, Zn, Cu, and Mg are a few examples of minerals whose mean concentration in the dry matter has decreased in a number of plant-based meals (Athar et al., 2020) [5] . Fortification of critical vitamins, antioxidants, and micronutrients has been emphasized in recent developments in conventional plant breeding. Adequate genetic variation in concentrations of β-carotene, other functional carotenoids, iron, zinc, and other minerals must exist among cultivars in order for conventional breeding to increase the micronutrient density of staple foods Dwivedi et al. (2023) [13] . This allows for the selection of nutritionally appropriate breeding materials.
Biofortification through Genetic Engineering
The direct use of biotechnology to alter an organism's genes is known as genetic engineering or genetic modification. By transferring desirable traits from one organism to another, genetic engineering makes it possible to create new cultivars that are elite and significantly increase their value (Athar et al., 2020) [5] . It has the potential to improve edible crops' nutritional content and bioavailability. Additionally, it presents special chances to enhance nutritional quality and provide additional health advantages. Numerous vegetable crops have undergone genetic modification to enhance characteristics like improved flavor or nutritional status, as well as to decrease bitterness, slow ripening, greater nutritional status, seedless fruit, increased sweetness, and anti-nutritional effects (Bouis and Saltzman, 2017) [7] .
Conclusion
By distributing naturally fortified foods to those with limited access to commercially promoted fortified foods, which are more easily accessible in metropolitan areas, biofortification offers a practical way to address malnourished populations in relatively remote rural locations. Therefore, commercial fortification and biofortification are quite complimentary. Adequate intakes of a variety of nutrients and other substances, in combinations and amounts that are yet unclear, are ultimately necessary for healthy nutrition. Therefore, increasing the consumption of a variety of nonstaple foods is the best and last way to end undernutrition as a public health issue in developing nations. To achieve this, though, it will need several decades, well-informed government policies, and a large investment in agricultural research as well as other public and on-farm infrastructure. In summary, according to M.S. Swaminathan, "GM foods have the potential to solve many of the world's hunger and malnutrition problems, and to help protect and preserve the environment by increasing yield, quality, and reducing reliance upon chemical pesticides." However, governments still face numerous obstacles, particularly in the fields of industrial policy, regulation, safety testing, and food labelling.
References
- Afify RRM, El-Nwehy SS. Nano fertilizers with algae extract as biostimulant affecting growth, bulb yield, and quality of onion. SABRAO J Breed Genet. 2023;55(6):2128-2139.
- Adeyeye SAO, Idowu-Adebayo F. Genetically modified and biofortified crops and food security in developing countries: A review. Nutr Food Sci. 2019;49(5):978-986.
- Awuchi CG, Igwe VS, Amagwula IO, Echeta CK. Health benefits of micronutrients (vitamins and minerals) and their associated deficiency diseases: A systematic review. Int J Food Sci. 2020;3(1):1-32.
- Atukunda P, Eide WB, Kardel KR, Iversen PO, Westerberg AC. Unlocking the potential for achievement of the UN Sustainable Development Goal 2 "Zero Hunger" in Africa: Targets, strategies, synergies and challenges. Food Nutr Res. 2021;65:29219.
- Athar T, Khan MK, Pandey A, Yilmaz FG, Gezgin S. Biofortification and the involved modern approaches. J Elementol. 2020;25(2):717-731.
- AOAC. Official Methods of Analysis. 20th ed. Washington (DC): Association of Official Agricultural Chemists; 2016.
- Bouis HE, Saltzman A. Improving nutrition through biofortification: A review. Glob Food Secur. 2017;12:49-58.
- Bouis HE, Saltzman A. Improving nutrition through biofortification: Evidence from HarvestPlus, 20032016. Glob Food Secur. 2017;12:49-58.
- Beebe S. Biofortification of common bean for higher iron concentration. Front Sustain Food Syst. 2020;4:573449.
- Choudhary RC, Kumaraswamy RV, Kumari S. Nanoenabled approaches for sustainable agriculture: A review. J Plant Nutr. 2020;43(10):1532-1548.
- Cichy K, Chiu C, Isaacs K, Glahn R. Dry bean biofortification with iron and zinc. In: Biofortification of Staple Crops. Singapore: Springer; 2022. p. 225-270.
- Das A, Laha S, Mandal S, et al. Preharvest biofortification of horticultural crops. Amsterdam: Elsevier Inc.; 2017.
- SL, Garcia-Oliveira AL, Govindaraj M, Ortiz R. Biofortification to avoid malnutrition in humans in a changing climate: Enhancing micronutrient bioavailability in seed, tuber, and storage roots. Front Plant Sci. 2023;14:1119148.
- Darshan SN, Vijaykumar NM, Vrashabh G, Vinay GC, Shankara MCM. Effect of varying levels of nitrogen on growth and yield of spinach (Spinacia oleracea). J Emerg Technol Innov Res. 2019;6(6):305-310.
- Dobosy P, Endredi A, Sandil S, Vetesi V, Záray G. Biofortification of potato and carrot with iodine by applying different soils and irrigation with iodinecontaining water. Front Plant Sci. 2020;11:593047.
- Gomathi M, Vethamoni PI, Gopinath P. Biofortification in vegetable crops -A review. Chem Sci Rev Lett. 2017;6:1227-1237.
- Garg M, Sharma N, Sharma S, Kapoor P, Arora P. Biofortified crops generated by breeding, agronomy, and transgenic approaches are improving lives worldwide. Front Nutr. 2018;1:1-33.
- Ikram NA, Ghaffar A, Khan AA, Nawaz F, Hussain A. Foliar iodine application: A strategy for tomato biofortification and yield optimization. J Plant Nutr. 2025;48(3):540-556.
- Jangra A, Tiwari S. Empowering nutrition: Biofortification to combat hidden hunger. In: Advances in Agri-Food Systems. Vol II. 2025. p. 285-305.
- Kumar A, Singh R, Yadav S. Nanotechnology-based delivery systems for enhancing crop productivity. Agric Syst. 2021;190:103089.
- Kalia P, Soi S, Muthukumar P. Marker-assisted introgression of the gene for enhancing β-carotene content in Indian cauliflower. In: IV Int Symp Mol Markers Hortic. 2017;1203:121-128.
- Martin AR, Broadley MR, Poblaciones MJ. Soil and foliar zinc biofortification of broccoli: Effects on plant growth and mineral accumulation. Crop Pasture Sci. 2020;71(5):484-490.
- Montañez R, Golob E, Xu S. Human cognition through the lens of social engineering cyber-attacks. Front Psychol. 2020;11:1755.
- Rahim FP, Rocio CG, Adalberto BM, Rosaura SCL, Maginot NH. Agronomic biofortification with selenium in tomato crops (Solanum lycopersicum L.). Agriculture. 2020;10(10):486-495.
- Roriz M, Carvalho SMP, Castro PML, Vasconcelos MW. Legume biofortification and the role of plant growth-promoting bacteria in sustainable agriculture. Agronomy. 2020;10(3):1-13.
- Saltzman A, Birol E, Oparinde A, Andersson MS, Asare-Marfo D, Gonzalez C, et al. Availability, production, and consumption of crops biofortified by plant breeding. Ann N Y Acad Sci. 2017;1390:104-114.
- Sharma P, Aggarwal P, Kaur A. Biofortification: A new approach to eradicate hidden hunger. Food Rev Int. 2017;33(1):1-21.
- Smoleń S, Barański R, Smoleń LI, Sady W. Combined biofortification of carrot with iodine and selenium. Food Chem. 2019;300:125202.
- Sharma S, Patel D, Singh R. Nano-encapsulated PGPR for chilli growth promotion. J Plant Growth Regul. 2020;39(3):785-795.
- Vandana T, Akhilesh S, Parveen S, Prabhat K, Shilpa. Biofortification of vegetable crops for vitamins, minerals and other quality traits. J Hortic Sci Biotechnol. 2022;28(10):1-13.
- O'Hare TJ, Fanning KJ, Martin IF. Zeaxanthin biofortification of sweet-corn and factors affecting zeaxanthin accumulation and colour change. Arch Biochem Biophys. 2015;572:184-187. doi:10.1016/j.abb.2015.01.015 American Chemical Society Publications doi:10.3389/fpls.2023.1119148 PMC
- Parida AK, et al. Impacts of salinity stress on crop plants: Improving salt tolerance through genetic and molecular dissection. Front Plant Sci. 2023;14:1241736. doi:10.3389/fpls.2023.1241736
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