Lead (Pb) severely impairs plant growth, yet the role of WRKY transcription factors in Pb tolerance in maize remains largely unknown. Here, we identified a Pb-responsive WRKY transcription factor, ZmWRKY4, whose transcript levels were rapidly and strongly induced in maize leaves following Pb exposure. Physiological and biochemical analyses showed that overexpression of ZmWRKY4 substantially enhanced Pb tolerance in maize. Transgenic lines exhibited significantly lower malondialdehyde (MDA) levels and reduced electrolyte leakage than wild-type plants. In addition, ZmWRKY4 overexpression increased catalase (CAT) activity and effectively limited H₂O₂ accumulation. Further analyses revealed that ZmWRKY4 positively regulates ZmCAT1, a key antioxidant gene involved in H₂O₂ scavenging, under Pb stress. Electrophoretic mobility shift assays and ChIP-qPCR collectively confirmed that ZmWRKY4 directly binds to W-box elements within the ZmCAT1 promoter in vivo and in vitro, thereby activating its transcription. Together, these findings define a previously uncharacterized ZmWRKY4-ZmCAT1 regulatory module that enhances antioxidant capacity and mitigates oxidative damage during Pb stress. This work provides new insights into the molecular mechanisms underlying heavy metal tolerance in maize and identifies a promising genetic target for developing Pb-resilient crop varieties.

NAC transcription factors are key regulators involved in diverse cellular processes, stress responses, and developmental pathways in plants. However, their roles in female cone development of Torreya grandis, a representative gymnosperm species, remain largely unexplored. In this study, we performed a comprehensive identification and analysis of NAC transcription factors in T. grandis to investigate their potential functions in female cone development. A total of 82 TgNAC members containing conserved NAM domains were identified, distributed unevenly across 11 chromosomes. Phylogenetic analysis with Arabidopsis NACs classified them into 15 groups, with TgNACs represented in 10 groups and showing a notable enrichment in the TERN clade on chromosome 2. Promoter cis-element analysis revealed correlations between regulatory elements and expression patterns. Tissue-specific expression profiling indicated clear functional specialization, with some TgNACs showing no detectable expression in the examined tissues. During female cone development, several TgNACs were highly expressed in the early stages, whereas TgNAC72, TgNAC76 and TgNAC82 were upregulated during the latter stages. Among these, TgNAC72 exhibited the highest overall expression level. Subcellular localization confirmed TgNAC72 is localized in the nucleus. Dual-luciferase assays further demonstrated that TgNAC72 activates the TgBGLU13 promoter, suggesting its role in starch and sucrose metabolism. Collectively, these findings provide novel insights into the regulatory involvement of TgNACs in reproductive organ development.