1 Introduction

Micronutrient deficiency, also known as “hidden hunger”, affects billions of people around the world. This situation will lead to poor attention, a high rate of illness, an increase in mortality rate, and also affect working ability (Welch and Graham, 2004; Kumar et al., 2019; Avnee et al., 2023). Potato is a staple crop that is widely consumed around the world. Especially in some developing countries where there are not many types of food and limited sources of nutrition, it has great potential in alleviating micronutrient deficiencies (Avnee et al., 2023; Seid et al., 2023). If the contents of vitamins and minerals in potatoes, such as iron, zinc, vitamins A, C and E, can be increased, the problems of these nutritional deficiencies can be effectively improved. It is not only beneficial to improving overall health, but also can reduce the social costs caused by malnutrition (Welch and Graham, 2004; Strobbe et al., 2018; Jiang et al., 2020).

Improving the nutritional quality of potatoes through breeding is a sustainable and low-cost method to deal with micronutrient deficiencies. Biofortification is a means to increase the content of vitamins and minerals in crops, which can be achieved through traditional breeding or genetic engineering. This method has achieved good results on many staple food crops (Saltzman et al., 2017; Strobbe et al., 2018; Kumar et al., 2019). Studies have already developed rice rich in beta-carotene and high-speed iron corn varieties (Welch and Graham, 2004; Strobbe et al., 2018). If the content of micronutrients in potatoes can be increased, it will help reduce “hidden hunger” in those areas where potatoes are the staple food (De Lepeleire et al., 2018; Seid et al., 2023). Nowadays, breeding techniques are becoming increasingly advanced, such as marker-assisted selection and metabolic engineering, which can help breed new potato varieties that have both nutrition and no impact on yield (Goyer, 2017; Strobbe et al., 2018; Seid et al., 2023).

This study reviewed the research progress of biofortification of essential vitamins and minerals in potatoes, and analyzed the application effects of different techniques such as traditional breeding and genetic engineering in improving the nutritional quality of potatoes. The research also pointed out some challenges currently faced in potato nutritional breeding, and at the same time summarized the available opportunities. This study proposes future research directions, aiming to provide reference and support for the cultivation and promotion of nutrient-rich potato varieties.

2 Current Status of Vitamin and Mineral Content in Potatoes

2.1 Levels of key vitamins in common potato varieties

Potatoes are an important source of vitamins, especially vitamin C and B vitamins. A 150-gram potato can provide approximately 45% of the recommended daily intake (RDA) of vitamin C, as well as 10% of vitamin B6, 8% of niacin and 6% of folic acid (Suttle, 2008). Nowadays, genetic engineering is also used to increase the vitamin content in potatoes. Some studies have shown that after genetically modifying potatoes to overexpress the PDX-II gene, the content of vitamin B6 increased by 107% to 150% (Bagri et al., 2018; Hameed et al., 2018). There are also significant differences in thiamine and folic acid contents among different potato varieties. The thiamine content of some genotypes is more than twice that of common varieties (such as Russet Burbank), and the folic acid content even reaches four times (Goyer and Sweek, 2011).

2.2 Mineral content in potatoes

Potatoes are also an important source of minerals. It can provide 18% of the daily required potassium, 6% of copper, phosphorus and magnesium, and approximately 2% of calcium and zinc (Pandey et al., 2023). Different potato varieties vary greatly in mineral content. The contents of phosphorus, potassium, sulfur and zinc in red-skinned potatoes are generally higher than those in brown-skinned potatoes, while the contents of calcium, magnesium and sodium in brown-skinned potatoes are higher (Pandey et al., 2023). At present, breeding research is also striving to increase the iron and zinc contents in potato tubers to enhance their nutritional value (Paget et al., 2014; Bradshaw, 2019).

2.3 Variation in vitamin and mineral levels among different potato cultivars

Different potato varieties have significant differences in the content of vitamins and minerals. Studies have found that some original cultivated varieties and wild potatoes have significantly higher levels of thiamine and folic acid than common cultivated varieties (Goyer and Sweek, 2011; Robinson et al., 2015). Some advanced breeding varieties also vary greatly in the mineral content of tubers. Except for iron, the differences in other minerals are quite significant. The mineral content of red-skinned potatoes is generally higher than that of brown-skinned potatoes and potato chip-specific varieties (Pandey et al., 2023).

3 Breeding Strategies for Enhancing Vitamin Content

3.1 Conventional breeding approaches for increasing vitamin content

Conventional breeding mainly increases the vitamin content in potatoes by selecting and hybridizing superior varieties. This type of breeding method mainly focuses on increasing the levels of vitamin B9 (folic acid) and vitamin C, and also attaches importance to the accumulation of carotenoids (such as lutein and zeaxanthin) in yellow and orange potatoes. Red and purple-fleshed potatoes have also become the focus of breeding attention because they are rich in anthocyanins and have antioxidant effects. Although there is rich genetic diversity in both current modern varieties and local varieties in the Andes region, rapid screening of these nutritional traits and ensuring the absorption effect (bioavailability) of these nutrients in the human body remain practical challenges (Bradshaw, 2019).

3.2 Marker-assisted selection (MAS) targeting genes associated with vitamin synthesis

Marker-assisted selection (MAS) is a method that combines molecular markers with traditional breeding techniques, which can screen out plants with ideal traits more efficiently and accurately. MAS has been successfully applied to select genes related to important agronomic traits such as disease resistance (Francia et al., 2005; Slater et al., 2017). MAS can also be used to identify and screen the key genes that control the synthesis of vitamin B6, vitamin C and carotenoids in terms of improving nutrition. One advantage of MAS is that it can screen out the target traits in the early stage of plant growth, which can greatly shorten the breeding cycle (Beketova et al., 2021; Brar et al., 2021) However, the effect of MAS is also affected by some factors, such as the genetic basis of the target trait, the degree of association between molecular markers and genes, and the number of samples available for analysis (Babu et al., 2004; Francia et al., 2005).

3.3 Use of transgenic techniques to enhance specific vitamin levels in potatoes

Genetic modification technology can increase the content of certain vitamins in potatoes by introducing specific genes. After introducing the PDX-II gene of Arabidopsis thaliana into potatoes, the content of vitamin B6 in tubers can be significantly increased, and at the same time, the tolerance of plants to abiotic stresses such as drought can also be enhanced (Bagri et al., 2018). Genetic engineering has also been used to increase the content of beta-carotene (A precursor of vitamin A) in potatoes, thereby cultivating the so-called “golden potatoes” (Bradshaw, 2019). Although these methods are very effective and can precisely regulate the target nutrients, they also face some challenges, such as the public's acceptance of genetically modified foods and policy approval restrictions (Hameed et al., 2018; Ahmad et al., 2022).

4 Breeding Strategies for Enhancing Mineral Content

4.1 Identifying and utilizing genotypes with high mineral accumulation

To increase the mineral content in potatoes, the first step is to identify the superior genotypes rich in minerals. Studies have shown that potato tubers have significant genetic differences in mineral content, and some varieties perform particularly well in nutrients such as potassium, calcium, magnesium and zinc (Lal et al., 2020; Pandey et al., 2023). Studies have found that the thiamine content of certain varieties is more than twice that of common varieties (such as Russet Burbank), and the folic acid content is also four times higher, which provides important genetic resources for nutrition-intensive breeding (Goyer and Sweek, 2011). When conducting genetic assessment on diploid potatoes, it was found that the heritability of trace elements such as iron and zinc was at a medium level, indicating that these traits could be effectively improved through breeding methods (Paget et al., 2014).

4.2 Developing biofortified potatoes using MAS and genomic tools

Marker-assisted selection (MAS) and genomic technology play an important role in the development of nutrition-fortified potatoes. These advanced methods can help researchers identify genes related to mineral content more accurately. For instance, through genome-wide association studies (GWAS), quantitative trait loci (QTL) related to the contents of potassium, manganese and zinc in potato tubers have been identified (Pandey et al., 2023). By using this genomic information, breeders can select parents with ideal traits more efficiently and accelerate the cultivation of new varieties. Combined with the prediction of genomic breeding values, it can also help screen out superior varieties that not only have high mineral content but also retain important agronomic traits such as high yield and disease resistance.

4.3 Role of root traits and mineral uptake efficiency in breeding for improved mineral content

In the breeding of increasing the mineral content in potatoes, root traits and mineral absorption efficiency play an important role. The process by which potatoes absorb minerals from the soil and transport them to the tubers is largely influenced by the structure and function of their root systems. Studies have pointed out that different potato varieties vary greatly in the mechanisms of mineral absorption and utilization, mainly due to different genotypes (Lal et al., 2020). By choosing those potato types with larger root systems and deeper distribution, breeders can cultivate varieties with higher mineral absorption efficiency. This method can not only enhance the nutritional level of tubers but also improve the utilization rate of nutrients, which is conducive to achieving a more sustainable way of potato production.

5 Genetic Basis of Vitamin and Mineral Accumulation

5.1 Key genes involved in vitamin synthesis pathways in potatoes

The genetic mechanism of vitamin synthesis in potatoes has been studied extensively, and scientists have identified several key genes. Overexpression of the PDX-II gene can significantly increase the content of vitamin B6 in tubers, while making plants more resistant to abiotic stresses such as salinity and oxidation (Bagri et al., 2018). By introducing the AT-HPT and At-γ-TMT genes of Arabidopsis thaliana, the content of α -tocopherol (vitamin E) in potatoes also increased significantly, and the stress tolerance was improved (Upadhyaya et al., 2020). In addition, researchers also significantly increased the folic acid level in tubers by simultaneously introducing multiple genes related to vitamin B9 (folic acid) synthesis, including GTPCHI, ADCS, HPPK/DHPS and FPGS. This provides a new idea for nutrition-enhanced breeding (Figure 1) (De Lepeleire et al., 2018).

5.2 Genes and loci associated with mineral transport and accumulation

The mineral content in potatoes is controlled by multiple genes and genetic loci, which regulate the transport and accumulation processes of minerals within the plant. Studies have found that the mineral content in potato tubers varies greatly and is related to multiple specific sites. The genome-wide association study (GWAS) revealed that quantitative trait loci (QTL) related to potassium and manganese content were found on chromosome 5, while zinc content was associated on chromosome 7 (Pandey et al., 2023). Studies on diploid potato populations also show that the traits of trace elements such as iron and zinc are mainly controlled by additive effects, and the heritability is at a medium level. This indicates that these nutrients can be effectively improved through breeding selection (Paget et al., 2014). There are also significant genetic differences in thiamine and folic acid contents among different varieties. This genetic diversity provides important resources for future nutrition-enhanced breeding (Goyer and Sweek, 2011).

5.3 Insights from genome-wide association studies (GWAS) and QTL mapping

Genome-wide association studies (GWAS) and quantitative trait locus (QTL) localization provide important clues for understanding the genetic basis of vitamin and mineral accumulation in potatoes. These methods can identify genomic regions related to the content of various nutrients in tubers. For instance, GWAS studies have found that there are QTLS related to potassium and manganese content on chromosome 5, and zinc content on chromosome 7, which is helpful for a better understanding of the genetic control mechanisms of these nutritional traits (Pandey et al., 2023). In diploid potatoes, QTL mapping also revealed a significant genetic correlation between mineral content and other tuber traits, providing breeders with a theoretical basis and technical framework for improving micronutrients (Paget et al., 2014).

6 Advances in Molecular Tools for Potato Breeding

6.1 CRISPR/Cas9 and gene-editing technologies for targeted trait improvement

CRISPR/Cas9 is a precise and efficient gene editing tool that has brought about significant breakthroughs in the field of genetic improvement and is also widely used in the breeding of crops such as potatoes (Huang, 2024). This technology can directly edit the genes of plants themselves without the need to introduce exogenous DNA, and thus non-genetically modified products can be obtained (Figure 2) (Chen et al., 2019; Ahmad et al., 2022). In potatoes, CRISPR/Cas9 has been used to enhance the nutritional quality of tubers, such as increasing the contents of vitamin C, β -carotene and methionine, while also reducing unfavorable components such as steroidal sugar alkaloids and acrylamide (Tussipkan and Manabayeva, 2021). With the disclosure of the whole genome sequence of potatoes and the optimization of the transformation system, the use of CRISPR/Cas9 has become more convenient, making it a powerful tool in potato nutritional breeding (Hameed et al., 2018; Tussipkan and Manabayeva, 2021; Das et al., 2023).

6.2 High-throughput phenotyping and genotyping for rapid selection

High-throughput phenotyping and genotyping techniques have become very important tools in modern potato breeding. High-throughput phenotyping technology can rapidly measure the traits of a large number of plants, such as growth rate, tuber size and disease resistance, without damaging the plants through advanced imaging and sensing equipment. High-throughput genotype technology can also rapidly identify genetic markers related to these traits, providing support for marker-assisted selection (MAS) and genomic selection (GS) (Chen et al., 2019; Ahmad et al., 2022). The combination of these two types of technologies has greatly enhanced the efficiency and accuracy of breeding, accelerated the cultivation of superior varieties, and also contributed to improving the nutritional quality and agronomic traits of potatoes.

6.3 Integrating transcriptomic and proteomic approaches to uncover regulatory networks

Combining transcriptomics and proteomics helps to gain a deeper understanding of the molecular regulatory mechanisms of potatoes during their development and stress response. Transcriptomics is a tool for studying RNA expression and can reveal the expression of genes under different conditions. Proteomics, on the other hand, focuses on the functions and changes of proteins. The combined use of these two methods can help researchers identify important genes and proteins related to key traits such as nutrient accumulation, disease resistance and stress tolerance, thereby gaining a more comprehensive understanding of the physiological mechanisms of potatoes. Scientists have utilized transcriptomic and proteomic analyses to identify candidate genes and proteins related to the increased content of vitamins and minerals in tubers, which provides important targets for nutritional breeding (Hameed et al., 2018; Ahmad et al., 2022).

7 Case Study: Breeding Potatoes for Higher Iron and Zinc Content

7.1 Background and importance of iron and zinc in potato breeding

Iron and zinc are essential trace elements for the human body. Iron is involved in oxygen transport and energy metabolism, while zinc is very important for the immune system and DNA synthesis. In many developing countries, deficiencies of iron and zinc are very common, which may lead to health problems such as anemia and low immunity (Andre et al., 2007; Seid et al., 2023). As an important staple food crop, potatoes have great potential in supplementing iron and zinc in the diet. Increasing the contents of iron and zinc in potatoes through breeding is helpful to alleviate micronutrient deficiencies and improve the health status of the population (Burgos et al., 2007; Bradshaw, 2019; Pandey et al., 2023).

7.2 Methods: screening high-iron and zinc genotypes and utilizing genetic modification

To cultivate potato varieties with higher iron and zinc content, researchers usually combine traditional breeding and genetic improvement techniques. A large number of varieties and breeding materials need to be screened and their iron and zinc contents evaluated. Studies have found that there are significant differences in iron and zinc concentrations among different potato genotypes, and some varieties in the Andean region have particularly high mineral contents (Andre et al., 2007; Burgos et al., 2007). Modern tools such as genome-wide association studies (GWAS) and genomic selection (GS) have been used to identify QTL loci related to iron and zinc content, helping breeders select parents more accurately (Pandey et al., 2023). Genetic engineering can also introduce key genes that control the absorption and storage of iron and zinc to increase the mineral levels in tubers. Some methods can also increase the bioavailability of iron and zinc by reducing anti-nutritional factors that hinder mineral absorption (Zimmermann and Hurrell, 2002; Bradshaw, 2019).

7.3 Results and lessons learned: successes, challenges, and future directions

Some achievements have been made in breeding work. Researchers have identified multiple potato genotypes rich in iron and zinc. In the breeding program of the International Potato Center, the iron content of some nutritionally fortified potato clones exceeded 32 mg/kg, and the zinc content exceeded 25 mg/kg (Sosa et al., 2018). In the field trials in Ethiopia, the study found that there were significant genetic differences in the contents of iron and zinc, and they had high heritability. This indicates that it is feasible to improve these traits through breeding (Seid et al., 2023). However, there are still some challenges. There is often a negative correlation between the content of iron and zinc and the output, which makes it difficult to improve both aspects simultaneously. Environmental conditions and the interaction between genotype and environment can also significantly affect the mineral content in tubers, increasing the complexity of the breeding process (Burgos et al., 2007; Nassar et al., 2012).

Whether potato varieties rich in iron and zinc can truly be effectively absorbed by the human body still requires further research. That is to say, merely increasing the content is not enough; attention should also be paid to the bioavailability of these minerals (Bradshaw, 2019). Future research should expand the genetic basis, for example, by incorporating diverse germplasm resources such as local varieties in the Andes region into breeding programs (Andre et al., 2007). To accelerate the screening efficiency, high-throughput phenotypic techniques such as X-ray fluorescence spectroscopy can be used to rapidly detect the mineral content of a large number of varieties (Sosa et al., 2018). Combining traditional breeding methods with modern genetic improvement techniques can also form a synergistic effect, which can not only increase the contents of iron and zinc, but also enhance their absorption rates (Zimmermann and Hurrell, 2002; Bradshaw, 2019).

8 Environmental and Agronomic Factors Influencing Vitamin and Mineral Content

8.1 Role of soil properties and fertilizers in affecting mineral levels in potatoes

Soil properties and fertilization methods are the key factors affecting the mineral content of potatoes. Studies have found that the use of organic fertilizers is more conducive to improving the nutritional components in potatoes than traditional chemical fertilizers. Organic fertilizers can usually increase the contents of phenolic substances, vitamin C and vitamin B9, and at the same time reduce the levels of harmful heavy metals such as cadmium (Cd) and nickel (Ni) (Rempelos et al., 2023). Spraying trace nutrients (such as boron, copper, iron, manganese, molybdenum and zinc) can also effectively increase the mineral content of potatoes. Studies show that this approach has increased the iron content in potatoes by 70% and the zinc content by 27%, significantly enhancing their nutritional value (Ierna et al., 2020). The planting location can also affect the mineral levels. For instance, the mineral content of potatoes grown in the Artova region of Turkey is significantly higher than that in other regions (Karan, 2023).

8.2 Impact of growing conditions on vitamin content

Growth conditions, such as light and water supply, can significantly affect the vitamin content in potatoes. Especially the climatic conditions during the planting season, such as rainfall, temperature and solar radiation, not only affect the yield, but also the quality of nutrition. These climatic factors can affect the synthesis of vitamin C and B9 in tubers, thereby influencing the nutritional level of the final harvest. Studies have found that climatic conditions have a greater impact on the yield and quality of potatoes than agronomic measures such as the previous crop or pest and disease management (Rempelos et al., 2023).

8.3 Interaction between genetic potential and environmental factors

The content of vitamins and minerals in potatoes is jointly influenced by the genetic potential of the variety and environmental conditions. Different potato varieties and clones perform differently in terms of nutrient accumulation due to their different genetic compositions. Studies have found that potatoes with cream-colored tubers perform best in terms of minerals such as potassium (K), phosphorus (P), magnesium (Mg), zinc (Zn), copper (Cu), and manganese (Mn), indicating that they have a genetic advantage in absorbing and storing these minerals. But the planting environment is also very important. For instance, in the study conducted by Tokat in Turkey, the mineral content of potatoes grown in different regions varied significantly, indicating that environmental conditions also have a significant impact on nutritional levels (Karan, 2023). Combining appropriate fertilization methods with the genetic potential of varieties can further improve the nutritional quality of tubers (Ierna et al., 2020).

9 Challenges and Limitations in Breeding for Enhanced Vitamin and Mineral Content

9.1 Trade-offs between yield, quality, and nutritional traits in breeding

In potato breeding, there are often certain contradictions and balance issues when it comes to simultaneously increasing the content of vitamins and minerals as well as other important traits (such as yield and quality). Increasing the concentrations of micronutrients such as iron and zinc may sometimes have adverse effects on tuber yield or other quality traits. Studies have shown that although there are significant genetic variations in nutrients such as iron and zinc, since these traits are often negatively correlated with yield, it is very challenging to improve the performance of both aspects simultaneously in breeding (Seid et al., 2023). In traditional breeding programs, more emphasis is often placed on disease resistance and processing performance, which may lead to the neglect of the goal of improving nutritional levels, resulting in difficulties in balancing multiple goals (Suttle, 2008).

9.2 Genetic diversity constraints in existing potato germplasm

The genetic diversity of potato germplasm resources is also one of the important challenges faced in breeding. Although different varieties and wild species show significant differences in vitamin and mineral contents, the genetic basis of some traits is still relatively narrow. Studies have found that different potato clones and species vary significantly in terms of thiamine and folic acid contents, but the genetic diversity of these nutritional traits is still limited overall (Goyer and Sweek, 2011; Robinson et al., 2015). This limitation makes it difficult for breeders to find parent materials that have both high nutritional content and good agronomic traits such as yield and resistance, thereby reducing the possibility of successful breeding (Paget et al., 2014).

9.3 Economic and technical challenges in developing and adopting improved varieties

In the process of promoting new varieties of potatoes rich in vitamins and minerals, there are still many economic and technical challenges. The commercialization of genetically modified potatoes is limited by consumers' acceptance and the restrictions of relevant regulations, which is a major obstacle in promotion (Hameed et al., 2018). At present, the lack of technology for rapid screening of certain nutrients also makes the breeding process more complicated (Bradshaw, 2019). Economic issues are equally important. The development cost of new varieties is relatively high, and the market’s acceptance of biofortified potatoes will also affect the promotion effect. If farmers think that the profit of new varieties is not high or the market demand is insufficient, they may not be willing to plant them (Lal et al., 2020).

10 Future Perspectives and Recommendations

10.1 Leveraging multi-omics approaches to accelerate trait discovery and breeding

The integration of multiple “omics” technologies such as genomics, transcriptomics, proteomics and ionomics provides a new approach for accelerating the discovery of important traits and improving the breeding efficiency of nutrition-fortified potatoes. These techniques can help researchers gain a comprehensive understanding of the genetic and molecular mechanisms behind nutrient accumulation and other traits in potato tubers. For instance, iomics can rapidly analyze the mineral composition in a large number of potato samples, providing important data support for biological enhancement targets such as iron and zinc enrichment (Carvalho and Vasconcelos, 2013). By combining multiple omics tools, scientists can identify key genes related to nutrient absorption, transport and accumulation, thereby promoting the development of new potato varieties with higher nutritional value (Carvalho and Vasconcelos, 2013; Pandey et al., 2023).

10.2 Combining traditional breeding with modern molecular tools for enhanced efficiency

The combination of traditional breeding with modern molecular tools such as CRISPR/Cas9 and TALENs can significantly enhance the efficiency of potato nutritional breeding. Traditional breeding has achieved certain results in improving nutrients such as iron, zinc and vitamins in tubers (Goyer and Sweek, 2011; Bradshaw, 2019). The precision and rapidity of modern molecular tools can make these breeding efforts more effective. For instance, CRISPR/Cas9 has been successfully used to develop new potato varieties without exogenous DNA, significantly increasing the contents of vitamin C and β -carotene (Hameed et al., 2018; Ahmad et al., 2022). By combining traditional methods with gene editing technology, breeders can cultivate new varieties that have both high nutritional value and conform to consumer preferences and policy standards more quickly (Hameed et al., 2018; Kumari et al., 2018).

10.3 Policy and collaborative efforts for global adoption of improved potato varieties

To promote the global dissemination of nutritionally fortified potatoes, the joint efforts of policy support and multi-party cooperation are needed. Collaboration among the government, scientific research institutions and the private sector is of great significance. The introduction of policies to encourage the development and promotion of biofortified crops is conducive to solving the global problem of micronutrient deficiencies (Robinson et al., 2015). Cooperation can also promote the sharing of genetic resources, technologies and experiences, and accelerate the breeding and promotion process of high-nutrition potato varieties. Carrying out public awareness about the health benefits of biofortified potatoes can also help increase consumer acceptance and market demand (Karan, 2023). Through improving policy mechanisms and strengthening international cooperation, the potential of biofortified potatoes in improving the global nutritional status will be better exerted (Robinson et al., 2015; Bradshaw, 2019).

Acknowledgments

The author extends gratitude to the two anonymous peer reviewers for their feedback on the manuscript of this study and to Ms. Xie for her assistance in organizing the data.

Conflict of Interest Disclosure

The author affirms 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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