Legume Genomics and Genetics 2025, Vol.16, No.2, 63-71 http://cropscipublisher.com/index.php/lgg 69 co-expression network, it is also possible to identify which gene modules they jointly respond to stress with. Although this approach cannot directly state "who regulates whom", it can help depict an overall picture. In contrast, proteomics is not as widely used, but it can bridge the gap between the transcriptional level and the protein level, verifying whether the WRKY protein is truly translated and participates in the reaction (Roorkiwal et al., 2020; Naqvi et al., 2024). 7.3 Gene editing tools (e.g., CRISPR/Cas) in WRKY functional analysis The emergence of CRISPR/Cas9 has brought the functional research of WRKY into an era of "precise operation". Although there haven't been many research cases of directly editing WRKY with CRISPR on chickpeas yet, the application of this technology in other crops is already quite mature, including regulating stress-resistant genes and improving agronomic traits, etc. If the editing advantages of CRISPR are combined with transcriptome or proteome data, it is theoretically possible to lock onto target genes more quickly and achieve targeted modification. That is to say, in the future, to improve the stress resistance of chickpeas, CRISPR may not be absent (Razzaq et al., 2021). 8 Future Directions and Applications of WRKYGenes in Chickpea Breeding 8.1 Potential of WRKYs in marker-assisted selection (MAS) Not all transcription factors are suitable as breeding markers, but WRKY is an exception, especially those that stand out in terms of disease resistance and stress tolerance. Like WRKY40, it was once positioned in important areas related to heat resistance and wilt disease resistance. These loci were not guessed out of thin air but were confirmed step by step through GWAS, QTL analysis and transcriptome data (Priyadarshini et al., 2023). All these pieces of evidence combined make WRKY a highly credible candidate for MAS. Under other environmental conditions such as salt stress, EST-SSR molecular markers related to WRKY have also been developed, which further indicates their usability in molecular breeding (Tarinejad et al., 2024). Of course, some members of the WRKY family may not be so "useful", which requires more meticulous screening. 8.2 Integration into stress-resilient chickpea breeding programs To truly utilize these genetic markers, a systematic breeding process is needed in coordination. Nowadays, multi-omics technology is developing rapidly, and many studies have begun to attempt to apply WRKY linkage markers to actual breeding. For instance, in high-temperature or severe wilt environments, previous studies have cultivated chickpea strains with better performance through marker-assisted backcrossing and genomic selection (Mohanty et al., 2024; Soorni et al., 2025). However, this plan is not universally applicable. The performance of some varieties in specific regions still needs to be verified. However, the overall trend is clear - WRKY is gradually transforming from a research subject in the laboratory to a practical tool that can be used in the field. 8.3 Challenges and prospects in translating genomics to field application Ultimately, transforming genomic data into actual agronomic traits is no easy task. Although the WRKY gene seems "versatile", their regulatory networks are often complex, involving multiple pathways. The upregulation of one gene may both enhance resistance and bring about negative growth effects. Moreover, the field environment itself is highly variable, and the interaction between genes and the environment can easily mask the original expression trend. In addition, the promotion of genetically modified crops is still subject to regulatory restrictions in many regions. However, with the emergence of CRISPR/Cas9, high-throughput phenotypic platforms, and more systematic field validation methods, these challenges are no longer insurmountable obstacles (Javed and Gao, 2023). With these technologies as support, the practical application prospects of the WRKY gene are actually closer to reality than ever before. Acknowledgments We would like to thank the anonymous reviewers for their detailed review of the draft. Their specific feedback helped us correct the logical loopholes in our arguments. 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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