## 1 Introduction  
The economic losses caused by plant-parasitic nematodes are estimated at USD 173 billion annually worldwide [1]. The rice white tip nematode (RWTN) _Aphelenchoides besseyi_ is a parasitic nematode that is widely present in crops, and more than 200 plant species can act as hosts for these nematodes. RWTN is widely distributed in almost all rice-growing areas in the world, causing losses of up to 50% in flooded rice [2]. It is difficult to implement a good means for thorough control of RWTN once it outbreaks. It is one of the top ten plant-parasitic nematodes and has been listed as a quarantined plant pest in many countries [3, 4]. Finding more effective and safe control measures is one of the main directions for research focusing on RWTNs and other nematodes. Unlike animals, plants lack specialised immune cells and rely exclusively on innate immunity to defend themselves against microbial invaders. The plant immune system operates on a sophisticated two-tier system, PAMP-triggered immunity (PTI) and effector-triggered immunity (ETI) [5]. As a dynamically changing three-dimensional network structure in eukaryotic cells, the actin cytoskeleton participates in the basic life activities of cells and facilitates a dynamic reorganisation process under the regulation of various actin-binding proteins (ABPs). In turn, the ABPs are involved in the perception and response of cells to external biotic and abiotic stimuli and balance and resist the threats of external stimuli to cell life activities [6]. In recent years, many studies have shown that the plant actin cytoskeleton is involved in the immune processes of plant PTI and ETI [6; 7; 8] and plays an important role in plant sensing and pathogen resistance [9]. In the plant-pathogen co-evolution process, pathogens use various strategies to avoid the plant immune response and maintain their virulence. At the same time, plants have evolved new defence strategies to resist the infection of pathogens. This \"Zig-Zag\" model is currently one of the many focus areas in plant-pathogen interaction research, and the discovery and analysis of its mechanisms will help us to better understand and control plant diseases [5]. Plant nematodes can secrete effectors to plant cells through nematode organs, such as the stylet and body wall. This affects the growth and metabolism of plants, dynamic balance of the cytoskeleton in plant cells, and defence responses of the plant. Consequently, these changes create a suitable environment for nematodes to complete their growth and development [10]. Therefore, studying the functions of these effectors and their interactions with hosts will help us to understand the nematode pathogenic mechanism and will allow for the selection of appropriate targets to control plant nematodes. Most studies have focused on the effectors expressed in the oesophageal glands of nematodes or the effectors secreted by their stylet [11]. These effectors are considered to be the key factors in the nematode parasitic process. For example, _Globodera rostochiensis_ SPRYSECE-19 and GrUBCEP12, _Meloidogyne incognita_ Mi-CRT, _Meloidogyne javanica_ MjTTL5, _Meloidogyne graminicola_ MgGPP, and _M. graminicola_ MgMO237 have been shown to inhibit plant defence responses in different ways [12; 13; 14; 15; 16; 17; 18]. The effector MiPFN3 of _M. incognita_ disrupts the actin cytoskeleton of plants to promote parasitism [19]. In addition, some studies have demonstrated that the effectors secreted by nematode body walls are crucial in the infection and pathogenicity of nematodes [20; 21]. For example, the peroxidase secreted by the hypodermis of _G. rostochiensis_ and _M. incognita_ can promote the development of nematodes by scavenging reactive oxygen species (ROS) in the host [22; 23]. Furthermore, the MiMIFs of _M. incognita_ secreted by the hypodermis can interact with _Arabidopsis thaliana_ annexin, suppressing plant immunity and promoting nematode parasitism ([PERSON] et al., 2019).