Journal of Energy Bioscience 2025, Vol.16, No.4, 205-215 http://bioscipublisher.com/index.php/jeb 212 comparative genomics studies have shown that C3 crops (such as wheat) themselves have the basis of C4 pathway genes. These genes have formed new tissue expression patterns during evolution, providing genetic resources for the engineering of C4 traits. In wheat, C4 enzymes such as PEPC, NADP-ME, MDH and PPDK have been identified and their activities have been detected at different growth stages and in tissues. The high expression of some genes is related to the improvement of photosynthetic efficiency. Wheat grains and panicles exhibit stronger C4 enzyme activity under stress such as drought, indicating that the C4 pathway plays a significant role in stress resistance and carbon utilization efficiency in non-lobar organs. With the development of synthetic biology and genetic engineering technology, progress has been made in the expression and localization of C4-related genes in C3 crops. However, to achieve a complete C4 pathway, it is still necessary to solve the problems of spatial division and the reconstruction of specific anatomical structures (such as Kranz structures). To achieve C4 wheat, multi-disciplinary collaboration is required, including molecular biology, genetics, plant physiology, synthetic biology and computational modeling. In-depth research on systems biology and gene regulatory networks can help break through the limitations of single enzyme engineering, promote multi-gene expression, tissue-specific regulation and anatomical structure remodeling. International cooperation projects (such as the C4 Rice Project) have provided references for this kind of cross-disciplinary cooperation. In the future, the C4 wheat project can be advanced in phases. The first step is to deeply analyze the expression regulation and function of C4-related genes in wheat; The second step is to achieve modular introduction of multiple genes and tissue-specific expression using synthetic biology methods. The third step is to explore and reshape the Kranz structure or a structure with similar functions; The fourth step is to optimize the C4 trait through field trials and molecular breeding to enhance yield and resource utilization efficiency. The ultimate goal is to cultivate C4 wheat with high light efficiency, strong stress resistance and sustainable development, providing feasible solutions for global food security and climate-resilient agriculture. Acknowledgments The authors sincerely appreciates the valuable opinions and suggestions provided by the three anonymous reviewers, whose meticulous review greatly helped me improve the quality of this 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. References Abramoff R., Ciais P., Zhu P., Hasegawa T., Wakatsuki H., and Makowski D., 2023, Adaptation strategies strongly reduce the future impacts of climate change on simulated crop yields, Earth's Future, 11(4): e2022EF003190. https://doi.org/10.1029/2022EF003190 Alenazi A., Bianconi M., Middlemiss E., Milenković V., Curran E., Sotelo G., Lundgren M., Nyirenda F., Pereira L., Christin P., Dunning L., and Osborne C., 2023, Leaf anatomy explains the strength of C4 activity within the grass species Alloteropsis semialata, Plant, Cell & Environment, 46(8): 2310-2322. https://doi.org/10.1111/pce.14607 Alvarez C., and Maurino V., 2023, Adaptive diversity in structure and function of C4 photosynthetic components, Biochemical Society Transactions, 51(3): 1157-1168. https://doi.org/10.1042/BST20221279 Asseng S., Martre P., Maiorano A., Rötter R., O'Leary G., Fitzgerald G., Girousse C., Motzo R., Giunta F., Babar M., Reynolds M., Kheir A., Thorburn P., Waha K., Ruane A., Aggarwal P., Ahmed M., Balkovič J., Basso B., Biernath C., Bindi M., Cammarano D., Challinor A., De Sanctis G., Dumont B., Rezaei E., Fereres E., Ferrise R., García-Vila M., Gayler S., Gao Y., Horan H., Hoogenboom G., Izaurralde R., Jabloun M., Jones C., Kassie B., Kersebaum K., Klein C., Koehler A., Liu B., Minoli S., Martin M., Müller C., Kumar N., Nendel C., Olesen J., Palosuo T., Porter J., Priesack E., Ripoche D., Semenov M., Stöckle C., Stratonovitch P., Streck T., Supit I., Tao F., Van Der Velde M., Wallach D., Wang E., Webber H., Wolf J., Xiao L., Zhang Z., Zhao Z., Zhu Y., and Ewert F., 2018, Climate change impact and adaptation for wheat protein, Global Change Biology, 25: 155-173. https://doi.org/10.1111/gcb.14481 Bachir D., Saeed I., Song Q., Linn T., Chen L., and Hu Y., 2017, Characterization and expression patterns of key C4 photosynthetic pathway genes in bread wheat (Triticum aestivum L.) under field conditions, Journal of Plant Physiology, 213: 87-97. https://doi.org/10.1016/j.jplph.2017.03.002
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