Journal of Energy Bioscience 2025, Vol.16, No.4, 205-215 http://bioscipublisher.com/index.php/jeb 211 crops (such as wheat) have decreased significantly in high temperatures and droughts. Moreover, the benefits of CO2 fertilization are limited and are easily offset by high temperatures (Asseng et al., 2018; Makowski et al., 2020; Pequeno et al., 2021; Rezaei et al., 2023). In contrast, C4 crops perform better under drought and high temperatures. If wheat can achieve C4 engineering, it will enhance its ability to cope with climate change, especially in low latitudes and arid regions (Rangan et al., 2016; Pequeno et al., 2021; Wang et al., 2022). At the same time, climate-smart measures (such as variety improvement and management optimization) can further reduce yield loss and even lead to net yield growth in some places (Pequeno et al., 2021; Sedebo et al., 2022; Abramoff et al., 2023). 7.3 Economic and social implications If the C4 project wheat is successfully promoted, it may bring about multiple economic and social benefits. Firstly, increased production and enhanced climate adaptability contribute to ensuring global food security, especially in developing countries that are highly threatened by climate change (Rangan et al., 2016; Pequeno et al., 2021; Martre et al., 2024). However, to reach the maximum yield potential, a large amount of nitrogen fertilizer input is required, which may increase environmental pressure and production costs (Jobe et al., 2020; Martre et al., 2024). Farmers adopting climate-smart measures (such as planting new varieties and optimizing management) can increase production by more than 34%, thereby increasing income and promoting rural economic development (Sedebo et al., 2022). In terms of policy-making, it is also necessary to consider the knowledge and practical experience of small-scale farmers to promote the popularization of technology and sustainable development. 8 Future Prospects and Research Priorities 8.1 Multi-institutional collaborations The C4 photosynthesis project is very complex and requires multi-institutional cooperation on a global scale. International projects like the C4 Rice Project have proved that cooperation plays an important role in resource integration, technology sharing and standard setting. In the future, if interdisciplinary and transnational cooperation can be promoted, breakthroughs in fields such as genomics, systems biology and synthetic biology can be accelerated, and the feasibility and efficiency of C4 engineering can be improved (Sage and Zhu, 2011; Schuler et al., 2016; Pradhan et al., 2022; Prasanna et al., 2025). 8.2 Bridging basic and applied research For the successful engineering of C4, it is essential to have a thorough understanding of the photosynthetic mechanisms, gene regulatory networks, and structures like the Kranz anatomy of C3 and C4. At present, basic research has identified many key genes and regulatory elements and studied their expression patterns. But more efforts are needed to transform these findings into engineering strategies that can be applied to wheat. In the future, priority should be given to closely integrating basic research with applied research, forming a complete innovation chain from molecular mechanisms to field performance (Matsuoka et al., 2003; Cui, 2021; Pradhan et al., 2022; Chen et al., 2023; Raturi et al., 2024). 8.3 Integrating C4 traits with other innovations Combining C4 traits with other innovative traits, such as stress resistance and efficient resource utilization, is an important direction for enhancing the adaptability and yield of wheat. The overexpression of the C4 gene not only enhances photosynthetic efficiency but also makes crops more drought-tolerant and heat-resistant. Future research can explore the combined effect of C4 traits with new technologies such as climate-smart crops and enhanced rhizosphere carbon sinks, to achieve comprehensive improvement of multiple traits (ansson et al., 2018; Yadav and Mishra, 2020; JMukundan et al., 2024; Talukder et al., 2024; Prasanna et al., 2025). 9 Conclusion In recent years, significant progress has been made in the research of C4 photosynthetic pathway engineering in wheat. Research has found that there are C4-related genes in the wheat genome, and the activity and expression of C4 enzymes can be detected in specific parts such as grains and panicles. These enzymes have a positive impact on carbon assimilation and yield under conditions such as high temperature and drought. Molecular evolution and
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