TGG_2025v16n4

Triticeae Genomics and Genetics 2025, Vol.16 http://cropscipublisher.com/index.php/tgg © 2025 CropSci Publisher, registered at the publishing platform that is operated by Sophia Publishing Group, founded in British Columbia of Canada. All Rights Reserved.

Triticeae Genomics and Genetics 2025, Vol.16 http://cropscipublisher.com/index.php/tgg © 2025 CropSci Publisher, registered at the publishing platform that is operated by Sophia Publishing Group, founded in British Columbia of Canada. All Rights Reserved. CropSci Publisher is an international Open Access publishing specializing in Triticeae genome, trait-controlling, Triticeae gene expression and regulation at the publishing platform that is operated by Sophia Publishing Group (SPG), founded in British Columbia of Canada Publisher CropSci Publisher Editedby Editorial Team of Triticeae Genomics and Genetics Email: edit@tgg.cropscipublisher.com Website: http://cropscipublisher.com/index.php/tgg Address: 11388 Stevenston Hwy, PO Box 96016, Richmond, V7A 5J5, British Columbia Canada Triticeae Genomics and Genetics (ISSN 1925-203X) is an open access, peer reviewed journal published online by CropSci Publisher. The journal publishes original papers involving in all aspects of Triticeae sciences. Subject areas covered comprise classical genetics analysis, structural and functional analysis of Triticeae genome, gene expression and regulation, efficient breeding of improved varieties, as well as transgenic varieties. It is positioned to meet the needs of breeders, geneticists, molecular biologists, and anyone, worldwide, engaged in the field of Triticeae research. All the articles published in Triticeae Genomics and Genetics are Open Access, and are distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. CropSci Publisher uses CrossCheck service to identify academic plagiarism through the world’s leading plagiarism prevention tool, iParadigms, and to protect the original authors’ copyrights.

Triticeae Genomics and Genetics (online), 2025, Vol. 16, No.4 ISSN 1925-203X http://cropscipublisher.com/index.php/tgg © 2025 CropSci Publisher, registered at the publishing platform that is operated by Sophia Publishing Group, founded in British Columbia of Canada. All Rights Reserved. Latest Content Cultivating High-Quality Wheat Varieties with High Nutritional Value Xian Zhang, Xuemei Liu Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 148-155 Effects of Irrigation Frequency on Dry Matter Accumulation and Water Use Efficiency of Wheat Yali Wang, Rugang Xu, Zhonghui He Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 156-165 Strategies to Improve Wheat's Drought and Heat Resistance ZhenLi Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 166-174 Identification of Key Transcription Factors Involved in Root Architecture of Barley Zhengqi Ma, Wei Wang Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 175-183 Molecular Breeding Strategies for Pyramiding Disease Resistance in Wheat Jin Wang, Xing Zhao, Fumin Gao Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 184-194

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 148-155 http://cropscipublisher.com/index.php/tgg 148 Feature Review Open Access Cultivating High-Quality Wheat Varieties with High Nutritional Value Xian Zhang, Xuemei Liu Hainan Provincial Key Laboratory of Crop Molecular Breeding, Sanya, 572025, Hainan, China Corresponding email: xuemei.liu@hitar.org Triticeae Genomics and Genetics, 2025, Vol.16, No.4 doi: 10.5376/tgg.2025.16.0016 Received: 09 May, 2025 Accepted: 19 Jun., 2025 Published: 07 Jul., 2025 Copyright © 2025 Zhang and Liu, This is an open access article published under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Preferred citation for this article: Zhang X., and Liu X.M., 2025, Cultivating high-quality wheat varieties with high nutritional value, Triticeae Genomics and Genetics, 16(4): 148-155 (doi: 10.5376/tgg.2025.16.0016) Abstract Wheat, as one of the most widely cultivated and consumed staple crops worldwide, plays a critical role in global food security and human nutrition. In this study, we explored comprehensive strategies for cultivating high-quality wheat varieties enriched with nutritional value, addressing the growing demand for healthier dietary options. Our approach integrates advances in genetic research-including gene mapping, transcriptomic profiling, and gene editing-with biofortification breeding methods and improved agronomic practices. We investigated genetic determinants for key traits such as protein, zinc, and iron content, and examined the application of GWAS and CRISPR technologies to enhance nutritional traits. Additionally, we assessed the impact of soil management, irrigation, and post-harvest practices on wheat quality. Case studies from India, Ethiopia, and China illustrate successful deployment of nutrient-rich wheat varieties and their implications for public health and food systems. Despite challenges such as yield-quality trade-offs and market constraints, the integration of multi-omics tools, precision agriculture, and global collaboration holds promise for future innovations. This study highlights the need for interdisciplinary efforts and policy support to ensure sustainable development of nutritionally superior wheat cultivars. Keywords Wheat biofortification; Nutritional quality; Genetic improvement; Agronomic practices; Precision agriculture 1 Introduction Wheat (Triticumspp.) is a very important food crop in the world. The staple food that many people eat every day, such as steamed bread and bread, is made of wheat. It can adapt to various climates and can be grown in many places, so the planting area is very large. According to research, wheat provides about half of the world's food calories (Khalid et al., 2023). Because wheat has high yield and strong adaptability, it is particularly important in some resource-poor areas (Hao et al., 2024). Wheat can be used to make a lot of things. Not only bread, but also noodles, pasta and other foods. So it accounts for a large part of the world's diet (Graziano et al., 2019). In addition to providing energy, wheat also contains a lot of nutrients. For example, protein, dietary fiber, and some important minerals such as iron, zinc and magnesium (Shamanin et al., 2024). However, the nutritional content of different types of wheat is not the same. Some old varieties, such as einkorn and emmer, contain more antioxidants and plant compounds than today's ordinary wheat, which may be more beneficial to the body (Mougiou et al., 2023). Wheat also contains gluten, which is critical to the quality of bread. But for some people, this ingredient can cause discomfort, such as celiac disease. So when breeding, in addition to considering yield and processing performance, we must also pay attention to its impact on health. The goal of this study is to find out how to breed more nutritious wheat varieties. We will study the genetic and environmental factors that affect wheat nutrition, and we will also see if traditional varieties and local varieties can help modern wheat improve nutrition. At the same time, we will analyze the biochemical composition and agronomic characteristics of wheat, hoping to find good genetic resources and breeding techniques to help breed nutritious and high-quality wheat varieties. Ultimately, these studies can provide references for future breeding work, allowing people to eat healthier and help improve food security.

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 148-155 http://cropscipublisher.com/index.php/tgg 149 2 Genetic Basis of Wheat Nutritional Quality 2.1 Genetic determinants of protein and micronutrient content The nutritional composition of wheat is determined by many genes. These genes affect the protein and various trace elements in the grain, such as iron, zinc and magnesium. The protein content (GPC) in wheat is one of the important indicators for judging whether it is nutritious. Studies have found that some gene loci (called SNPs) are distributed on chromosomes 1D, 3A, 3B, 3D, 4B and 5A, which are related to protein synthesis and nutrient transport processes (Kartseva et al., 2023). In addition, genes are also involved in wheat's absorption of macronutrients such as nitrogen, phosphorus and potassium. For example, SNPs related to these nutrients have also been found on chromosomes 1A, 1B, 1D and A3 (Aljabri and El-Soda, 2024). This information is very helpful for breeding, especially when you want to improve wheat nutrition. 2.2 Biofortification methods in wheat breeding Nowadays, biofortification is a widely used breeding method. Its goal is to select wheat varieties that contain more iron, zinc, and magnesium. Through these varieties, we can cultivate wheat with higher nutrition (Rabieyan et al., 2023; Petrović et al., 2024). Different varieties of wheat vary greatly in nutrient content. These differences can be used for selection during breeding. Not only common wheat varieties, but also some wild wheat and local old varieties are nutritious, and their genetic resources are very valuable. Introducing these excellent genes into modern varieties can improve the overall nutritional level of wheat (Zeibig et al., 2024). This method is particularly useful in some developing countries because many people rely on wheat as their staple food, but their nutritional intake is insufficient. 2.3 Genomics and transcriptomics tools for quality improvement Now that technology is becoming more and more advanced, there are more tools for improving wheat. Technologies such as "Genome-wide Association Study" (GWAS) and "Quantitative Trait Loci Location" (QTL) can help us find genes related to protein content and mineral accumulation (Figure 1) (Lou et al., 2020; Fradgley et al., 2022). With this data, breeders can more accurately select the desired varieties. Tools such as SNP chips and transcriptome analysis can also tell us which genes play a role in the synthesis and transport of nutrients (Khalid et al., 2023). These technologies can make breeding work more targeted and efficient, and help to improve the nutritional quality of wheat to a new level. 3 Agronomic Measures to Improve Wheat Quality 3.1 The role of soil and fertilization in nutrient absorption For wheat to absorb more nutrients, the soil must be good and the fertilization must be reasonable. Especially nitrogen fertilizer, it has a great impact on the protein in the grain and the overall quality. If the amount of nitrogen fertilizer and the time of fertilization can be arranged well, it will not only make the wheat grow better, but also reduce pollution (Liu et al., 2018; Melash and Ábrahám, 2022). In addition to chemical fertilizers, organic fertilizers and crop rotation are also useful. Using organic fertilizers can improve soil structure, make the soil healthier, and nutrients are more easily absorbed. Crop rotation can also help restore soil nutrients, thereby improving wheat yield and nutritional quality (Li et al., 2023). 3.2 Impact of irrigation and crop management Water is important for wheat, and planting methods can also affect its yield and quality. In arid areas, if water can be applied once and for all at the right time, coupled with deep loosening of the soil and proper use of fertilizers, wheat leaves will be healthier and photosynthesis will be stronger, so that the yield will naturally increase (Huang et al., 2024). In addition, the planting method can also be adjusted, such as appropriately changing the density of sowing. This can help wheat make better use of rainwater, especially in places where rainfall is irregular. This practice allows wheat to use water when it needs it, is less susceptible to drought, grows stronger, and has a more stable yield (Yang et al., 2021). In the final analysis, these methods are all about making water and fertilizer use more accurately and truly use them when and where wheat needs them most.

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 148-155 http://cropscipublisher.com/index.php/tgg 150 3.3 Impact of post-harvest handling and storage After wheat is harvested, how it is handled will also affect its quality. If the temperature and humidity can be controlled well, the wheat will not spoil easily and the nutrients can be preserved. However, if it is not handled properly, such as when the soil is compacted too tightly during harvesting, or the ground is too lumpy, it will not only affect the quality of the wheat, but may also make the soil hard, affecting the next season's planting (Al-Shammary et al., 2023). Therefore, after harvesting wheat, it is best to adopt some methods to protect the soil. For example, light tillage, try not to disturb the soil structure, and add scientific storage methods, which can help the wheat retain nutrients and make the quality more stable. Figure 1 Relationships among traits and the proportion of the trait heritability explained by QTL (Adopted from Fradgley et al., 2022) Image caption: a: Network analysis of all analysed milling, baking and micronutrients traits across two trial years identify eight distinct groups. Blue and red connecting lines indicate positive and negative correlations, respectively and line width is proportional to correlation strength. Only correlations with p < 0.001 are shown. b: Proportion of phenotypic variation explained by the broad sense heritability as well as all QTL included in a full model for SNP and haplotype-based analysis for meta-analysis across two trial years for all traits. GPC=Grain protein content (%); SPW specific weight (kg hl-1), SKCS single kernel characterisation system hardness, ER extraction rate (%); L*=Flour whiteness (Tristimulus L*); b*=Flour yellowness (Tristimulus b*); L*-b*=overall flour colour (Tristimulus L*-b*); HFN=Hagberg Falling Number (s); GA=MARVIN grain area (mm-1); GL=MARVIN grain length (mm); GW=MARVIN grain width (mm); TGW=MARVIN thousand grain weight (g); BWAP=DoughLab Bandwidth at Peak; DT=DoughLab Development time (s); MTI=DoughLab mixing tolerance index; PE=DoughLab Peak Energy; SO DoughLab Softening; ST=DoughLab Stability; WA=DoughLab Water absorption (%); SDS=SDS Sedimentation (ml); whole=mineral concentration in whole meal flour (mg/kg); white=mineral concentration in refined white flour (mg/kg) (Adopted from Fradgley et al., 2022)

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 148-155 http://cropscipublisher.com/index.php/tgg 151 4 Cultivate High-Quality and High-Nutrition Wheat Varieties 4.1 Integration of quality and yield traits When breeding, we should not only look at how much grain wheat can produce, but also whether it is nutritious. In the past, many breeding projects pursued high yields, but nutrition was easily overlooked. Now research is paying more and more attention to both aspects. Scientists hope to select varieties that are both high-yield and nutritious, such as those with high protein content and trace elements such as iron and zinc (Voss-Fels et al., 2019; Hao et al., 2022). Some traditional varieties and wild wheat contain some useful genes that can improve nutrition or yield. Introducing these genes into modern wheat can make the new varieties more nutritious while ensuring that the yield does not decrease. This approach is very helpful in improving people's malnutrition problems. 4.2 Hybrid vigor of hybrid wheat and its quality traits Hybrid wheat is to "match" two different wheat varieties to combine their respective advantages. The wheat bred in this way can not only have high yield, but also have better quality, such as more protein and stronger disease resistance (Zhao et al., 2015). But to breed good hybrid varieties, the most important thing is to choose the right "parents". Current genetic technology can help us analyze the effects of different combinations and improve the success rate. This method can select the most suitable combination and help cultivate high-quality wheat to meet people's demand for nutritious food. 4.3 Stress-resistant varieties with high-quality traits In recent years, climate change has become more and more obvious. Extreme weather such as droughts and heat waves are increasing. Therefore, we need wheat that can grow well in this environment. This kind of wheat must not only be drought-resistant and heat-resistant, but also maintain good quality. Now breeders have added these "stress-resistant" genes to high-quality wheat (Mondal et al., 2016). They use some new technologies, such as genomic selection and high-throughput phenotyping technology (Paux et al., 2022), to find suitable breeding materials more quickly. The wheat bred in this way is not only nutritious, but also can be grown in different regions. They help improve food security and better cope with complex environments in the future. 5 Case Study: Regional Implementation of Fortified Wheat Varieties 5.1 Success in India in promoting zinc biofortified wheat India has introduced zinc-enriched wheat varieties to reduce zinc deficiency. Zinc deficiency is a common health problem in India and neighboring countries. Farmers have helped wheat absorb more zinc by changing the way they apply fertilizers, such as using phosphorus-rich compost and zinc sulfate (Table 1). This method is called "biofortification". Studies have shown that it does increase the zinc content of wheat grains (Paramesh et al., 2020). This method not only makes wheat more nutritious, but also makes the soil healthier and the crops grow better. Therefore, it also helps promote more environmentally friendly agriculture (Sharma et al., 2019). Table 1 Influence of integrated P management and Zn application on P concentration and uptake by wheat (average of 2 years) (Adopted from Paramesh et al., 2020) Treatments Grain P concentration (%) Straw P concentration (%) Grain P (kg/ha) Straw P (kg/ha) Total P (kg/ha) Phosphorus levels Without P 0.28c 0.13b 10.4c 7.6c 17.9d P100-F 0.3 b 0.13b 13.5b 8.2c 21.7c P100-PEC 0.38 a 0.15a 18.1a 10.5b 28.6a P50-PEC+P50-F 0.37 a 0.15a 18.0a 11.0a 29.0a P75-PEC+VAM+PSB 0.38 a 0.15 a 17.3a 10.2b 27.4b Application of ZnSO4.7H2O Without Zn 0.34b 0.15a 13.9c 9.1b 23.0d 25kg-Soil 0.33b 0.13c 15.6b 9.1b 24.7c Two foliar ** 0.36a 0.15a 15.4b 9.8a 25.2b Soil+Foliar * 0.36a 0.14b 16.9a 10.0a 26.9a Note: **-Two foliar spray at anthesis and one week after anthesis stage; *-one foliar spray at one week after anthesis stage. Similar letter in a column indicates non-significance difference between treatments (Adopted from Paramesh et al., 2020)

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 148-155 http://cropscipublisher.com/index.php/tgg 152 5.2 Ethiopia uses high-protein wheat in food aid programs Ethiopia has introduced high-protein wheat in food aid programs to address malnutrition issues. This is particularly suitable for areas where wheat is the staple food. High-protein wheat provides more essential amino acids for the body, which helps improve nutrition. The local government selected varieties with high protein content and distributed this more nutritious wheat to those in need, such as children and pregnant women (Tanin et al., 2024). This example also shows that when breeding, we should not only focus on yield, but also pay attention to nutritional content, especially in areas in need of assistance. 5.3 China's innovation in wheat nutrigenomics China has also made many new attempts in wheat breeding, with the goal of making wheat both nutritious and high-yielding. Researchers have proposed a method called "High Nutrient Use Efficiency Fertilization" (High NUFER). This method can make wheat have high yield and more protein, reduce fertilizer use, and have less impact on the environment (Hou et al., 2023). They developed the most suitable fertilization plan based on the genes and actual growth of wheat to help improve the nutritional level. These explorations in China show that as long as the method is appropriate, it is possible to cultivate high-yield and nutritious wheat, and it can also be more environmentally friendly, which will have a positive impact on food security. 6 Challenges and Opportunities 6.1 Genetic and physiological trade-offs It is actually quite difficult to breed wheat that is both high-yielding and nutritious. Sometimes, in order to make wheat drought-resistant or disease-resistant, we may end up reducing its yield or affecting its quality. For example, some drought-resistant wheat may not produce much grain even though it is suitable for dry places. This is because these traits are not determined by a single gene, but by many genes and the environment (Bapela et al., 2022). Wheat genes are inherently complex. If you want it to be high-yielding, nutritious, disease-resistant and drought-resistant, it will be even more difficult (Mondal et al., 2016). Therefore, when breeding, you cannot just focus on one aspect. If you improve a certain trait, you may make others worse, so you have to try to balance it. 6.2 Socioeconomic and market constraints In addition to technical issues, there are also some social and market difficulties. Some people do not approve of nutritionally fortified wheat, especially genetically modified wheat. When promoting it, you may encounter people who do not understand it and the policy does not keep up (Bhalla, 2006). The current market demand for wheat is more about how well it can be processed, such as how well it is ground into flour and how well it makes bread. On the contrary, whether it is nutritious is not the main consideration. This also makes it difficult to promote many varieties that focus on "nutrition" (Subedi et al., 2023). In addition, breeding and promoting new varieties cost a lot of money, which is a big problem for research on nutritious wheat (Saquee et al., 2024). To solve these problems, on the one hand, more publicity is needed to make people aware of the benefits of nutritious wheat; on the other hand, the government should also introduce policies and provide some financial and technical support. 6.3 Policy and research gap The new technologies used in breeding, such as "genomic selection" and "marker-assisted breeding", are not widely used in practice. One reason is that policies cannot keep up and support is not strong enough (Paux et al., 2022). This affects the promotion speed of new technologies. In addition, whether fortified wheat is good for people if eaten for a long time has not been thoroughly studied. There are still many blanks in its impact on health and food security (Sharma et al., 2023). To solve these problems, scientists, governments and enterprises need to work together. Only through more communication and cooperation can we build a good mechanism to promote truly nutritious and promising wheat varieties so that more people can benefit from them. 7 Future Outlook 7.1 Integrating multi-omics technologies for nutritional breeding Now, more and more scientists are beginning to use "multi-omics" to improve the nutrition of wheat. These methods include genome, transcriptome, proteome and metabolome. Through these technologies, we can

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 148-155 http://cropscipublisher.com/index.php/tgg 153 understand more clearly how wheat nutrition is formed, and we can also find out the key genes that affect nutrient accumulation and stress resistance (Li et al., 2021). With this information, breeders can select new varieties more directional. Such wheat is not only high in protein, but also rich in various trace elements. At the same time, its yield will not decrease, and its resistance will not deteriorate (Katamadze et al., 2023). These new tools can also help us breed wheat suitable for different regions to better meet people's nutritional needs. 7.2 Precision agriculture helps improve quality Precision agriculture has been very popular in recent years. It uses some new technologies, such as remote sensing, soil monitoring and precision fertilization, to help farmers farm more scientifically. These technologies can make water and fertilizer use more accurately, avoid waste, and improve the quality of wheat (Yadav and Pyare, 2024). It can also monitor the growth of wheat at any time, such as whether there is a lack of water, fertilizer, or pests and diseases. If problems are found, farmers can take immediate measures to reduce losses and increase yields (Voss-Fels et al., 2019). If these precision agricultural technologies are combined with breeding technologies, high-yield and nutritious wheat can be grown, and damage to the environment can be reduced, making agriculture greener. 7.3 Global cooperation in wheat nutrition improvement Breeding nutritious and resilient wheat is not something that one or two countries can do alone. This requires global cooperation. Countries can share seed resources, breeding techniques and research results. Only in this way can we breed wheat varieties suitable for planting in different places (Rajaram, 2001). Some international projects are already doing this. For example, BREEDWHEAT is a good example. It brings together scientists and breeders to study the nutrition and yield issues of wheat (Paux et al., 2022). Through such cooperation, scientists can breed nutritious and high-quality wheat varieties more quickly. These varieties can also be more easily promoted to different countries, which is very helpful for improving global food security and human health. Acknowledgments We are grateful to Mrs.Guo for critically reading the manuscript and providing helpful comments that improved the clarity of the text. 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. References Aljabri M., and El-Soda M., 2024, Genome-wide association mapping of macronutrient mineral accumulation in wheat (Triticum aestivum L.) grain, Plants, 13(24): 3472. https://doi.org/10.3390/plants13243472 Al-Shammary A., Al-Shihmani L., Caballero-Calvo A., and Fernández-Gálvez J., 2023, Impact of agronomic practices on physical surface crusts and some soil technical attributes of two winter wheat fields in southern Iraq, Journal of Soils and Sediments, 23: 3917-3936. https://doi.org/10.1007/s11368-023-03585-w Bapela T., Shimelis H., Tsilo T., and Mathew I., 2022, Genetic improvement of wheat for drought tolerance: progress, challenges and opportunities, Plants, 11(10): 1331. https://doi.org/10.3390/plants11101331 Bhalla P., 2006, Genetic engineering of wheat--current challenges and opportunities, Trends in Biotechnology, 24(7): 305-311. https://doi.org/10.1016/J.TIBTECH.2006.04.008 Fradgley N., Gardner K., Kerton M., Swarbreck S., and Bentley A., 2022, Trade-offs in the genetic control of functional and nutritional quality traits in UK winter wheat, Heredity, 128: 420-433. https://doi.org/10.1038/s41437-022-00503-7 Graziano S., Marando S., Prandi B., Boukid F., Marmiroli N., Francia E., Pecchioni N., Sforza S., Visioli G., and Gullì M., 2019, Technological quality and nutritional value of two durum wheat varieties depend on both genetic and environmental factors, Journal of Agricultural and Food Chemistry, 67(8): 2384-2395. https://doi.org/10.1021/acs.jafc.8b06621 Hao B., Ma J., Si L., Jiang L., Wang X., Yao C., Ma S., Li C., Gao Z., and Wang Z., 2022, Did wheat breeding simultaneously alter grain concentrations of macro- and micro-nutrient over the past 80 years of cultivar releasing in China? Frontiers in Plant Science, 13: 872781. https://doi.org/10.3389/fpls.2022.872781

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Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 156-165 http://cropscipublisher.com/index.php/tgg 156 Review Article Open Access Effects of Irrigation Frequency on Dry Matter Accumulation and Water Use Efficiency of Wheat Yali Wang, Rugang Xu, Zhonghui He Modern Agricultural Research Center, Cuixi Academy of Biotechnology, Zhuji, 311800, Zhejiang, China Corresponding email: zhonghui.he@cuixi.org Triticeae Genomics and Genetics, 2025, Vol.16, No.4 doi: 10.5376/tgg.2025.16.0017 Received: 21 May, 2025 Accepted: 03 Jul., 2025 Published: 21 Jul., 2025 Copyright © 2025 Wang et al., This is an open access article published under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Preferred citation for this article: Wang Y.L., Xu R.G., and He Z.H., 2025, Effects of irrigation frequency on dry matter accumulation and water use efficiency of wheat, Triticeae Genomics and Genetics, 16(4): 156-165 (doi: 10.5376/tgg.2025.16.0017) Abstract Wheat (Triticum aestivumL.) is a globally essential cereal crop whose productivity is closely linked to water availability, particularly in water-limited regions. This study explores the effects of different irrigation frequencies on dry matter accumulation and water use efficiency (WUE) in wheat cultivation. We examined the physiological basis of biomass accumulation and analyzed how irrigation intervals influence partitioning among organs and developmental stage-specific responses. Further, we evaluated WUE in relation to irrigation frequency, considering agronomic implications and the interplay of root development, leaf structure, and molecular signaling pathways. A case study from a semi-arid wheat-growing region provided field-based insights into the impacts of irrigation frequency on yield, soil health, and practical outcomes. Our analysis highlights the trade-offs between water input and biomass productivity, emphasizing the importance of optimized irrigation scheduling. We conclude that moderate irrigation intervals can enhance WUE without severely compromising yield, though outcomes depend on local climate and soil conditions. Future research should focus on site-specific strategies using precision agriculture to improve sustainability under climate variability. Keywords Wheat; Irrigation frequency; Dry matter accumulation; Water use efficiency; Semi-arid agriculture 1 Introduction Wheat is one of the most important food crops in the world. It is critical to food security and agricultural development. In recent years, the increase in wheat yields is mainly due to variety improvement and the rational use of resources such as water and nitrogen (Hao et al., 2023; Ma et al., 2024). Among these resources, water plays the greatest role. Water supply directly affects the growth process of wheat, and also affects the accumulation of dry matter and the final yield. Reasonable irrigation can keep wheat photosynthesis strong and prolong the filling time, thereby improving the efficiency of water and nitrogen use (Li et al., 2018; Li et al., 2019; Lyu et al., 2020). The time and frequency of irrigation have a great influence on the distribution of dry matter and also affect the reuse of carbon reserves. These factors are closely related to grain development and final yield (Wang et al., 2011; Huang et al., 2014). In water-scarce areas, how to arrange the irrigation time becomes particularly important. Only in this way can we achieve both increased production and resource conservation (Wang et al., 2023). This study mainly aims to summarize recent research results on the effects of irrigation frequency on wheat dry matter accumulation and water use efficiency. We will pay special attention to several common irrigation methods, such as drip irrigation, micro-spraying and supplementary irrigation. How do these methods affect wheat physiological processes, dry matter distribution and resource utilization efficiency under different wheat varieties and different environments? By integrating research under different ecological conditions, we hope to provide some practical suggestions for future irrigation management to make wheat cultivation more productive, environmentally friendly and sustainable. 2 Dry Matter Accumulation in Wheat under Varying Irrigation Frequencies 2.1 Physiological basis of dry matter accumulation Wheat's dry matter is mainly produced through photosynthesis. The quality of photosynthesis is closely related to the availability of water and nutrients. Irrigation can help wheat maintain a large leaf area and fast photosynthesis,

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 156-165 http://cropscipublisher.com/index.php/tgg 157 and it can also slow down plant aging, thereby accelerating the accumulation of dry matter and allowing it to accumulate for a longer time, especially during the important stages of wheat growth (Li et al., 2023). Generally speaking, dry matter accumulation grows in an "S-shaped" pattern. If there is not enough water, wheat will reach the peak of accumulation earlier, but the rate and total amount of accumulation will be reduced (Yan et al., 2022). In addition, the process of transferring dry matter from the nutritional organs such as stems and leaves to the grains is also critical, and this process is also affected by irrigation and nitrogen fertilizer application (Xue et al., 2006). 2.2 Impacts of irrigation frequency on biomass partitioning The frequency of irrigation affects the distribution of dry matter in the wheat plant, such as to the stem, leaves, or ears. More frequent irrigation or more accurate timing of irrigation results in more dry matter being distributed to the ears and grains, which is important for increasing yield. For example, wheat that was irrigated four times during an important developmental stage had more total dry matter and more dry matter distributed to the ears than wheat that was irrigated only once or twice (Pal et al., 2000). When drip or sprinkler irrigation is used, more dry matter is distributed to the leaves and ears, and less to the stem and leaf sheaths, especially during the grain filling period. If water is insufficient, wheat will transfer previously stored nutrients to the grains to make up for the current loss of photosynthesis. 2.3 Developmental stage-specific responses When to irrigate is also important, depending on the developmental stage of the wheat. Irrigation during the jointing, booting, flowering and filling stages can prolong the accumulation time of dry matter and allow more dry matter to be transferred to the grain (Ma et al., 2024). However, if there is a lack of water during the tillering stage or milky stage, the total amount of dry matter and the proportion transferred to the ear will decrease significantly, and the final yield will be greatly affected (Han et al., 2022). Supplemental irrigation from the booting stage to the filling stage can not only increase the dry matter in the nutritional organs and grains, but also reduce the dependence on pre-flowering reserves. Different irrigation times have different effects on wheat of different varieties and in different environments, which will affect yield and water use efficiency together (Moradi et al., 2022). 3 Water Use Efficiency (WUE) in Response to Irrigation Frequency 3.1 Definition and agronomic importance of WUE Water use efficiency (WUE) of wheat refers to the amount of wheat yield or dry matter produced for each unit of water used. The higher the WUE, the more valuable the water is. This indicator is particularly important in water-scarce areas. Because water resources are scarce, improving WUE can help farmers save water while maintaining stable yields (Li et al., 2019b; You et al., 2022). 3.2 Effects of irrigation intervals on WUE How to arrange irrigation and the frequency of irrigation will directly affect WUE. Appropriate irrigation time, such as irrigation at the jointing stage and heading stage, can make wheat grow well and reduce unnecessary water waste, thereby increasing yield and WUE at the same time (Si et al., 2020). For example, in semi-humid areas, if 60 mm of water is applied at the jointing stage and heading stage respectively, relatively high WUE and yield can be obtained (Bian et al., 2016). Increasing the number of irrigations but watering less each time, such as using drip irrigation or micro-spraying, can also improve WUE. This can keep the soil moist and photosynthesis more sustained. However, if irrigation is too frequent or too much water is used, it may waste water, which will reduce WUE and may not necessarily increase yield (Figure 1). In the case of water-saving planting, even if the total yield is slightly reduced, WUE may be higher. This shows that a good balance must be found between yield and water saving (Stallmann et al., 2020). 3.3 Factors influencing WUE dynamics WUE is affected by many factors. For example, drip irrigation and micro-spraying combined with a reasonable irrigation frequency can improve WUE more than traditional irrigation methods, especially when water is insufficient (Hao et al., 2023). Soil type is also important. For example, loam or sandy soil, combined with a low

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 156-165 http://cropscipublisher.com/index.php/tgg 158 precipitation environment, is more likely to achieve higher WUE under water-saving conditions (Yu et al., 2020). Different wheat varieties have different water use efficiencies. In addition, the effect of irrigation is different at different growth stages, such as jointing, booting or filling. Another method called "active water" or "oxygenated water" has also been found to improve WUE, especially when water is scarce (Wang et al., 2022). However, it should be noted that although increasing the irrigation frequency can improve WUE, using advanced irrigation systems also requires investment. In the end, it depends on whether it is worth it and whether a good balance can be found between increasing yield and saving water (Fang et al., 2018). Figure 1 (A) Aboveground dry mass and (B) applied water use efficiency of wheat plants subjected to well-watering (ctr), continuous (cd) and pulsed (pd) drought, harvested at two time points (T1=77 d and T2=93 d after sowing). The applied water use efficiency was calculated for each pot as the ratio of aboveground plant dry mass to the cumulative amount of water received until harvest. At T2, values are given for vegetative (leaves and stems) and generative plant parts (ears). The boxes represent the interquartile ranges, whiskers extend to the 10% and 90% percentiles, respectively; solid lines show the medians, dashed lines the means. Outliers are shown as circles; when there was a significant effect of irrigation treatment, manual contrasts between selected groups were calculated and p values are given; ***p < 0.001; **p < 0.01; *p < 0.05; (n.s.) marginally significant (p < 0.1); n.s. not significant; n=10 (Adopted from Stallmann et al., 2020)

Triticeae Genomics and Genetics, 2025, Vol.16, No.4, 156-165 http://cropscipublisher.com/index.php/tgg 159 4 Mechanistic Insights and Physiological Responses 4.1 Root development and water uptake efficiency Wheat roots are responsible for absorbing water and are a key part of water use. The role of the root system is particularly evident under different irrigation frequencies and drought conditions. Drought-tolerant wheat varieties usually grow thicker and larger roots. These roots can grow and absorb water better, and can also activate genes related to carbon metabolism and hormone signaling, thereby enhancing drought resistance (Hu et al., 2018). Root shape and structure, such as lateral root length and number of root tips, are directly related to water absorption efficiency. Using molybdenum fertilizer can make roots grow better, such as making roots more permeable to water, and can also increase the expression of water channel proteins, thereby enhancing water absorption capacity. The method and time of irrigation can also affect root distribution. If irrigation is more frequent, there will be more roots in the upper soil layer and the ability to absorb water will also be stronger (Jha et al., 2017). In addition, wheat with long root hairs and heavy root sheaths can also absorb water more easily and help control water evaporation. 4.2 Leaf morphology and transpiration The coordination between leaves and roots also affects how water is used, especially during droughts or when there is little water. Sometimes wheat grows thinner roots to improve water absorption efficiency. Stable isotopes in leaves can be used to determine the current water status and transpiration rate of the plant (Brunel-Saldias et al., 2020). Studies have found that using molybdenum fertilizer can reduce leaf transpiration and allow roots to absorb more water, indicating that the direction of water flow has changed, and water is more concentrated in the roots for use rather than evaporating from the leaves (Wu et al., 2019). In addition, the relationship between some substances secreted by the roots and the surrounding microorganisms will also affect soil water retention, and these factors will also change the transpiration process (Rabbi et al., 2021). 4.3 Hormonal and molecular signals Plants use hormones to regulate their response to water. One of the most important is abscisic acid (ABA). When the soil dries out, the roots sense it and produce ABA. This hormone travels to the leaves, closing the stomata, which reduces water loss and helps the plant save water during later droughts (Saradadevi et al., 2017). Drought-tolerant wheat generally has stronger protective mechanisms, such as stronger antioxidant capacity and more active hormones related to root growth (Hu et al., 2024). There is also a substance called nitric oxide, which is also involved in regulating root development and enhancing water absorption during drought, and these regulations may also be affected by the trace element molybdenum (Wu et al., 2019). From a molecular perspective, some transcriptome and metabolome studies have found that drought-tolerant wheat activates some special pathways, such as synthesizing flavonoids, osmoprotectants, and enhancing energy metabolism. These changes can help roots grow better and cope with drought. 5 Agronomic and Environmental Considerations 5.1 Yield stability under different irrigation schemes Many studies have shown that wheat yield and water use efficiency (WUE) can be kept stable or even improved if irrigation time is chosen appropriately and water use is well controlled (Gao et al., 2022). This method will be even more effective if variety improvement is combined. Even in years with significant climate change, timely irrigation before sowing, during jointing or flowering can keep yields on track and reduce yield differences between years. In some places where water is scarce, appropriate reduction in water use (called "deficit irrigation") can increase WUE, and will have little impact on yield if soil conditions are suitable (Yu et al., 2020). In addition, combining scientific fertilization with water-saving irrigation can further improve yield stability and resource utilization efficiency (Huang et al., 2024). 5.2 Soil health and sustainability To ensure sustained high wheat yields, the soil must be kept healthy. Reducing irrigation and applying fertilizers properly are effective methods. For example, controlling the amount of water and combining it with appropriate nitrogen fertilizers can make the soil healthier, reduce nitrogen loss, and reduce pollution to the environment (Xu

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