# The Structure, Classification, Functional Diversity and Regulatory Mechanism of Plant C2H2 Transcription Factors

## TL;DR

This review summarizes the structure, function, and regulation of C2H2 transcription factors in plants, highlighting their roles in development and stress responses.

## Contribution

The paper provides a comprehensive overview of C2H2 TFs, including their classification, evolution, and regulatory mechanisms in plant stress adaptation.

## Key findings

- C2H2 TFs regulate floral organ development and pollen fertility in plants.
- They coordinate stress responses by modulating hormone signaling and reactive oxygen species homeostasis.
- Evolutionary differences between woody and herbaceous plants in C2H2 TFs are identified.

## Abstract

Cys2/His2-type zinc finger transcription factors (C2H2 TFs) constitute one of the largest transcription factor families in plants and play critical regulatory roles in growth, development, and stress responses. This review systematically summarizes the structural characteristics, classification systems, and evolutionary dynamics of this family, with a focus on their functional mechanisms in flower development, abiotic stress responses (drought, salt, low temperature, aluminum toxicity and so on), and biotic stress responses. We elucidate the molecular networks through which C2H2 TFs integrate hormone signals such as ABA to regulate reactive oxygen species homeostasis and stomatal movement. Through cross-species comparative analysis, we examine the evolutionary differences between woody and herbaceous plants and discuss current research limitations as well as future application prospects based on gene editing and multi-omics technologies. This review provides a theoretical reference for crop stress resistance breeding and quality improvement.

Cys2/His2-type zinc finger transcription factors (C2H2 TFs) constitute one of the largest and most functionally diverse transcription factor families in plants, playing core regulatory roles in multiple aspects of plant growth, development, and stress adaptation. Based on literature data from databases including PubMed (1995–April 2026) and integrated with bioinformatics analyses, this review provides a comprehensive overview of this family. We first summarize the structural characteristics and classification systems of C2H2 TFs, and elucidate their evolutionary dynamics from lower plants to angiosperms. Regarding their impact on plant organ development, beyond key biological processes, this review details the molecular mechanisms of C2H2 TFs in floral organ morphogenesis (e.g., petal, sepal, stamen, and ovule development), pollen fertility maintenance, and flowering time regulation. Concurrently, we systematically analyze their functional pathways in responses to abiotic stresses (drought, high salinity, low temperature, aluminum toxicity, etc.) and biotic stresses (pathogens, pests), clarifying the molecular networks through which they coordinate reactive oxygen species (ROS) homeostasis, stomatal movement, and osmotic regulation by modulating hormone signaling pathways such as ABA, SA, and JA. Furthermore, this review discusses major limitations of current research, including knowledge gaps concerning functional redundancy, pseudogenization phenomena, and cell type-specific regulation. We also provide perspectives on future research directions leveraging cutting-edge technologies such as CRISPR gene editing, single-cell sequencing, and multi-omics integration, as well as their application prospects in crop stress resistance breeding and quality improvement. This review provides ideas for in-depth research on the regulatory network and related functions of C2H2 TFs, and offers reference value for improving plant traits, enhancing plant resistance, and increasing the production of plant secondary metabolites.