1 Introduction

Micronutrient deficiencies, often termed “hidden hunger,” remain a pervasive global health challenge, particularly in regions where diets are dominated by a limited range of staple crops. Maize, as one of the world’s most widely cultivated and consumed cereals, provides a substantial portion of daily calories for millions, especially in sub-Saharan Africa, Latin America, and South Asia. However, its nutritional monotony—characterized by low levels of essential micronutrients such as vitamin A, iron, and zinc—contributes to widespread deficiencies and associated health burdens, including impaired growth, anemia, and increased susceptibility to disease (Nuss and Tanumihardjo, 2010; Galani et al., 2020; Goredema-Matongera et al., 2021; Msungu et al., 2022).

Given its central role as both a food and feed crop, maize presents a unique opportunity for nutritional enhancement. Its global reach and adaptability make it an ideal candidate for interventions aimed at improving dietary quality and combating malnutrition at scale (Nuss and Tanumihardjo, 2010; Palacios-Rojas et al., 2020). Biofortification, the process of increasing the nutrient content of crops through breeding, agronomic practices, or biotechnology, has emerged as a promising, cost-effective, and sustainable strategy. Unlike conventional fortification or supplementation, biofortification delivers nutritional benefits directly through staple foods, ensuring wide and long-term impact, particularly among rural and resource-poor populations (Garg et al., 2018; Galani et al., 2020; Goredema-Matongera et al., 2021; Msungu et al., 2022).

The advantages of biofortification are manifold: it is low-cost, requires minimal changes to existing food habits, and offers a sustainable solution to micronutrient deficiencies by leveraging the natural food system. Successful examples include the development and commercialization of maize varieties enriched with provitamin A, lysine, tryptophan, zinc, and iron, which have demonstrated improved health outcomes in target populations (Garg et al., 2018; Goredema-Matongera et al., 2021; Msungu et al., 2022).

This study will comprehensively review research progress in maize biofortification, exploring key strategies including conventional breeding, molecular techniques, and agronomic interventions. It will also review recent achievements and analyze challenges and future directions in achieving nutritional security. This study aims to provide researchers, policymakers, and practitioners with a deeper understanding of the importance and potential of maize biofortification in addressing global micronutrient deficiencies.

2 Nutritional Composition of Maize and Targeted Fortification Directions

2.1 Overview of maize nutritional content

Maize is a staple crop providing significant calories globally, with its kernels primarily composed of starch (about 70%~75%), moderate protein (8%~10%), and low fat (3%~5%). It also contains essential vitamins (notably B vitamins) and minerals (such as magnesium, zinc, and iron), though the levels of some micronutrients are often insufficient to meet daily requirements, especially in populations relying heavily on maize as a dietary staple (Nuss and Tanumihardjo, 2010; Galani et al., 2020; Palacios-Rojas et al., 2020).

Conventional maize varieties are limited by low concentrations of key nutrients: lysine and tryptophan (essential amino acids), provitamin A (β-carotene), iron, and zinc. These deficiencies contribute to widespread malnutrition, particularly in regions where maize is a primary food source (Nuss and Tanumihardjo, 2010; Galani et al., 2020; Palacios-Rojas et al., 2020).

2.2 Key targets for nutritional fortification

Quality Protein Maize (QPM) varieties have been developed to increase lysine and tryptophan content, addressing the poor protein quality of traditional maize. QPM can significantly improve the nutritional value of maize-based diets, especially for vulnerable populations (Nuss and Tanumihardjo, 2010; Palacios-Rojas et al., 2020).

Biofortification efforts have successfully increased provitamin A (β-carotene) content in maize, offering a sustainable solution to vitamin A deficiency. Provitamin A-biofortified maize varieties can provide a substantial portion of daily vitamin A requirements, especially in sub-Saharan Africa and other regions with high deficiency rates (Cabrera-Soto et al., 2018; Palacios-Rojas et al., 2020; Aman, 2021; Msungu et al., 2022).

Recent breakthroughs have enabled the development of maize varieties with significantly higher iron and zinc concentrations. For example, the identification of the ZmNAC78 gene has allowed for the doubling of iron content in maize kernels without yield penalties, while both genetic and agronomic approaches have improved zinc content and bioavailability (Maqbool and Beshir, 2018; Ahmad et al., 2023; Sahu, 2024).

Maize also contains phytochemicals such as anthocyanins and carotenoids, which have antioxidant properties and potential health benefits. Breeding programs are exploring ways to enhance these compounds to further improve the functional and nutritional quality of maize (Palacios-Rojas et al., 2020).

3 Major Strategies for Maize Biofortification

3.1 Traditional breeding strategies

Hybrid breeding and backcrossing have been foundational in introducing high-nutrient genes into elite maize lines, resulting in the release of hundreds of biofortified varieties enriched with iron, zinc, and provitamin A across diverse environments (Garg et al., 2018; Virk et al., 2021). Marker-assisted selection (MAS) has significantly improved breeding efficiency by enabling precise tracking of nutrient-related genes, thereby accelerating the development of varieties such as Quality Protein Maize (QPM) and provitamin A-rich maize (Maqbool et al., 2018). However, the expression of nutritional traits is often influenced by genotype × environment interactions, necessitating multi-environment testing and the use of breeding-friendly quantitative trait loci (QTLs) validated across genetic backgrounds to ensure trait stability and effectiveness (Maqbool et al., 2018; Sethi et al., 2023; Fatihah, 2024).

3.2 Genetic engineering and gene editing technologies

Transgenic approaches have enabled the introduction of exogenous nutrient metabolic pathways, exemplified by the development of high β-carotene (provitamin A) maize through the insertion of genes from other species. CRISPR/Cas-mediated gene editing has further advanced the field by allowing precise modification of endogenous genes controlling nutrient accumulation, such as crtRB1 and lcyE, which are key regulators of carotenoid biosynthesis (Maqbool et al., 2018). Multi-gene stacking and pathway optimization strategies are being explored to simultaneously enhance multiple nutrients and optimize metabolic flux, offering the potential for multi-nutrient biofortified maize varieties (Labuschagne, 2023; Sen et al., 2024).

3.3 Omics-driven and systems breeding

The integration of genomic, transcriptomic, and metabolomic data has revolutionized the dissection of nutrition-related networks in maize, enabling the identification of candidate genes and regulatory elements underlying complex traits (Sahito et al., 2024). Precision phenotyping platforms, combined with high-throughput omics data, have enhanced screening efficiency and facilitated the rapid selection of superior genotypes with improved nutritional profiles (Farooqi et al., 2022; Mahmood et al., 2022). These systems-level approaches support the mainstreaming of biofortification traits into competitive maize varieties and hybrids, ensuring sustainable impact at scale (Virk et al., 2021; Gedil et al., 2024).

4 Case Studies and Representative Achievements

4.1 Promotion and impact of quality protein maize (QPM)

QPM was developed by harnessing the opaque2 (o2) gene, which increases lysine and tryptophan content, overcoming the inherent protein quality limitations of conventional maize. Marker-assisted selection and backcrossing have been instrumental in incorporating the o2 gene into elite lines, resulting in QPM varieties with improved nutritional profiles (Prasanna et al., 2020; Goredema-Matongera et al., 2021).

QPM has been widely adopted in sub-Saharan Africa and Latin America, where it has demonstrated significant nutritional benefits, particularly for children and vulnerable populations. Field studies show that QPM hybrids not only enhance protein quality but also maintain competitive yields and agronomic performance under diverse environmental conditions (Prasanna et al., 2020; Goredema-Matongera et al., 2021).

4.2 Vitamin a-fortified maize

Vitamin A deficiency is a major public health issue in many maize-dependent regions. Biofortified maize varieties have been developed by introducing or selecting for favorable alleles of β-carotene synthesis genes such as psy1 and crtRB1, leading to increased provitamin A (β-carotene) content in maize kernels (Prasanna et al., 2020; Msungu et al., 2022).

Both transgenic and non-transgenic approaches have been used to enhance provitamin A in maize. Non-transgenic methods, such as marker-assisted selection for crtRB1 and lcyE alleles, have been widely adopted due to regulatory and consumer acceptance advantages, while transgenic strategies offer the potential for even higher provitamin A accumulation (Prasanna et al., 2020; Msungu et al., 2022).

4.3 Breeding progress in micronutrient (iron, zinc) enhancement

Significant progress has been made in developing and deploying maize varieties with elevated iron and zinc content. Field trials and multinational pilot programs have identified and released cultivars with improved micronutrient concentrations, contributing to better nutritional outcomes in target populations (Dhakal et al., 2022; Prasanna et al., 2020; Ahmad et al., 2023; Xue et al., 2023).

High β-carotene maize can help prevent vitamin A deficiency, while high-zinc maize contributes to improved micronutrient intake. By combining conventional breeding with marker-assisted selection (MAS) techniques, multiple highly adaptable cultivars have been developed in different countries according to local climate and agronomic conditions (Figure 1) (Prasanna et al., 2020). Genetic studies have identified key quantitative trait loci (QTLs) and candidate genes responsible for iron and zinc accumulation in maize kernels. These include loci involved in mineral uptake, transport, and deposition, providing valuable targets for marker-assisted breeding and further genetic improvement (Basnet and Khanal, 2022).

Figure 1 Provitamin A-enriched and high-Zn maize cultivars developed using conventional and molecular marker-assisted breeding and released for commercial cultivation in sub-Saharan Africa, Asia, and Latin America (Adopted from Prasanna et al., 2020)

5 Socioeconomic Impacts and Evaluation of Biofortified Maize Products

5.1 Nutritional impact assessment

Clinical trials and field surveys consistently demonstrate that biofortified maize can significantly improve nutritional status in target populations. For example, a randomized controlled trial in rural Malawi showed that consumption of selenium-biofortified maize flour led to substantial increases in serum selenium concentrations among women and children, providing strong evidence that agronomic biofortification can effectively address micronutrient deficiencies in real-world settings (Joy et al., 2022). Meta-analyses of quality protein maize (QPM) interventions reveal that replacing conventional maize with QPM results in a 12% increase in weight gain and a 9% increase in height among infants and young children with mild to moderate undernutrition, highlighting the positive impact on child growth in maize-dependent communities (Gunaratna et al., 2010). Additionally, studies in sub-Saharan Africa and South Africa report that provitamin A and zinc-biofortified maize varieties have the potential to alleviate vitamin A and zinc deficiencies, especially among children, with high acceptability and improved nutritional profiles (Siwela et al., 2020; Chawafambira et al., 2021; Goredema-Matongera et al., 2021; Msungu et al., 2022). Sensory evaluations indicate that women and children in rural Ethiopia accept QPM for complementary feeding, suggesting that consumer acceptance is unlikely to impede the nutritional impact of biofortified maize (Gunaratna et al., 2016).

5 Socioeconomic Impacts and Evaluation of Biofortified Maize Products

5.1 Nutritional impact assessment

Clinical trials and field surveys consistently demonstrate that biofortified maize can significantly improve nutritional status in target populations. For example, a randomized controlled trial in rural Malawi showed that consumption of selenium-biofortified maize flour led to substantial increases in serum selenium concentrations among women and children, providing strong evidence that agronomic biofortification can effectively address micronutrient deficiencies in real-world settings (Joy et al., 2022). Meta-analyses of quality protein maize (QPM) interventions reveal that replacing conventional maize with QPM results in a 12% increase in weight gain and a 9% increase in height among infants and young children with mild to moderate undernutrition, highlighting the positive impact on child growth in maize-dependent communities (Gunaratna et al., 2010). Additionally, studies in sub-Saharan Africa and South Africa report that provitamin A and zinc-biofortified maize varieties have the potential to alleviate vitamin A and zinc deficiencies, especially among children, with high acceptability and improved nutritional profiles (Siwela et al., 2020; Chawafambira et al., 2021; Goredema-Matongera et al., 2021; Msungu et al., 2022). Sensory evaluations indicate that women and children in rural Ethiopia accept QPM for complementary feeding, suggesting that consumer acceptance is unlikely to impede the nutritional impact of biofortified maize (Gunaratna et al., 2016).

5.2 Economic and social benefits analysis

Biofortification of maize contributes to reducing public health costs by addressing micronutrient deficiencies in populations with limited access to dietary supplements and fortified foods, particularly in low- and middle-income countries. By improving nutritional status, biofortified maize can help decrease the prevalence of deficiency-related diseases, potentially lowering healthcare expenditures and productivity losses (Goredema-Matongera et al., 2021; Msungu et al., 2022). Economic analyses indicate that the adoption of biofortified maize can enhance farmers’ income, especially when extension strategies effectively communicate both agronomic and nutritional benefits. In East Africa, farmer participation in extension programs and awareness of nutritional advantages were key drivers of QPM adoption, which in turn can improve household food security and livelihoods (De Groote et al., 2016; Jada and Van Den Berg, 2024). However, the economic impact may vary by context; for instance, in the Indian poultry sector, the cost-saving potential of biofortified maize as animal feed was found to be marginal due to the availability of alternative protein sources and limited awareness among small-scale producers (Krishna et al., 2014). Social factors, such as nutrition education and community health integration, further support the adoption and sustained impact of biofortified maize (Siwela et al., 2020; Jada and Van Den Berg, 2024).

6 Challenges in the Promotion and Application of Maize Biofortification

6.1 Technical and breeding bottlenecks

Achieving and maintaining genetic stability of nutritional traits, such as enhanced levels of zinc, provitamin A, or quality protein, remains a significant challenge. Many nutritional traits are controlled by multiple genes with small effects, making them difficult to improve and stabilize through conventional breeding. Environmental factors, such as soil nutrient availability and climate stress, can also affect the expression and consistency of these traits across different regions and seasons, complicating the adaptability of biofortified varieties (Lung’aho et al., 2011; Gupta et al., 2015; Maqbool and Beshir, 2018; Goredema-Matongera et al., 2021).

Transgenic and gene-edited maize varieties face stringent regulatory and safety evaluation requirements. These include comprehensive assessments of potential environmental impacts, food safety, and allergenicity, which can delay the release and adoption of new biofortified cultivars. Regulatory hurdles are particularly pronounced for genetically modified organisms (GMOs), often resulting in slow dissemination and limited public sector investment in transgenic approaches (Gupta et al., 2015; Garg et al., 2018).

6.2 Public acceptance and market promotion issues

Public awareness of the health benefits of biofortified maize is often low, especially in rural and resource-poor communities. Misconceptions about the safety and taste of biofortified products, particularly those developed through genetic modification, can hinder acceptance. Consumer education and outreach are critical to improving perception and encouraging adoption (Gupta et al., 2015; Palacios-Rojas et al., 2020; Msungu et al., 2022).

Biofortified maize varieties must compete with traditional, well-established varieties that may be preferred for their yield, taste, or processing qualities. Limited market incentives and lack of clear price premiums for biofortified products further discourage farmers and traders from adopting and promoting these varieties (Gupta et al., 2015; Palacios-Rojas et al., 2020; Goredema-Matongera et al., 2021).

6.3 International cooperation on resource and technology sharing

International collaboration is essential for accessing diverse germplasm and advanced breeding technologies. However, challenges such as intellectual property rights, benefit-sharing agreements, and regulatory differences between countries can restrict the free exchange of genetic resources and knowledge (Gupta et al., 2015; Goredema-Matongera et al., 2021).

There is an urgent need to establish robust international cooperation mechanisms to facilitate resource and technology sharing, harmonize regulatory frameworks, and support capacity building in developing regions. Such cooperation is vital for scaling up biofortification efforts and ensuring that the benefits reach vulnerable populations globally (Gupta et al., 2015; Goredema-Matongera et al., 2021).

7 Future Trends in Maize Biofortification

7.1 Multi-nutrient co-fortification strategies

Recent research is shifting from single-nutrient enhancement to the simultaneous improvement of multiple micronutrients in maize, such as provitamin A, zinc, iron, lysine, and tryptophan. This integrated approach addresses the complex nutritional deficiencies prevalent in maize-dependent regions, especially in sub-Saharan Africa. Developing maize varieties with multinutritional attributes is seen as a sustainable and cost-effective solution, with ongoing efforts to combine traits for improved adaptation to local stress conditions and dietary needs (Palacios-Rojas et al., 2020; Goredema-Matongera et al., 2021; Labuschagne, 2023). Advances in breeding and agronomic practices, including the use of plant growth-promoting rhizobacteria, are also contributing to the enrichment of several nutrients at once (Rajendran and Veeramani, 2022; Ahmad et al., 2023).

7.2 Frontier applications of gene editing and synthetic biology

Gene editing technologies, such as CRISPR-Cas, and synthetic biology are at the forefront of precision breeding for maize biofortification. These tools enable targeted modifications to enhance specific nutritional traits and facilitate the smart design of maize with tailored nutrient profiles. The integration of gene editing with conventional and marker-assisted breeding is expected to accelerate the development of multi-nutrient and climate-resilient maize varieties. Regulatory trends suggest that gene-edited crops may face fewer barriers than traditional GMOs, potentially speeding up their adoption and impact (Labuschagne, 2023).

7.3 Smart agriculture and precision biofortification pathways

The future of maize biofortification is closely linked to the adoption of smart agriculture technologies. Intelligent breeding models, supported by digital agriculture, genomic selection, and efficient nutrient monitoring, are being developed to optimize nutrient content and crop performance. Precision biofortification pathways leverage data-driven approaches to select and manage varieties with superior nutritional profiles, ensuring that biofortified maize meets both agronomic and nutritional targets efficiently (Rajendran and Veeramani, 2022; Labuschagne, 2023). These innovations are expected to enhance the scalability and sustainability of biofortification programs.

8 Concluding Remarks

Maize biofortification stands out as a promising and sustainable approach to address “hidden hunger”—the widespread micronutrient deficiencies affecting millions in maize-dependent regions. By leveraging both traditional breeding and modern biotechnological innovations, significant progress has been made in enhancing the nutritional quality of maize, including increased levels of provitamin A, zinc, iron, and essential amino acids. These advances have demonstrated measurable improvements in human health, such as reduced rates of vitamin A deficiency and iron-deficiency anemia, particularly among vulnerable populations in sub-Saharan Africa and other developing regions.

The integration of agronomic practices, genetic improvement, and microbial biofortification has further expanded the potential of maize to deliver essential nutrients efficiently and cost-effectively. Clinical and field studies confirm that biofortified maize varieties can significantly improve nutritional status and yield without compromising agronomic performance. However, the success of biofortification programs depends on coordinated efforts across scientific research, supportive policy frameworks, and active public engagement to ensure widespread adoption and sustained impact.

A sustainable biofortification strategy requires ongoing investment in research, robust extension services, and effective communication to raise awareness and acceptance among farmers and consumers. International collaboration and resource sharing are also critical to accelerate progress and ensure that the benefits of biofortified maize reach those most in need. Ultimately, maize biofortification offers a multifaceted solution to global nutritional challenges, with the potential to improve health, livelihoods, and food security for future generations.

Acknowledgments

Thank you to the anonymous peer review for providing targeted revision suggestions for the manuscript.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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