摘要
      消毒技术，特别是基于光的消毒，已经经历了巨大的发展和创新，但这种处理方法可以使细菌进入存活但不可培养的状态。由于其强大的耐受性，VBNC细菌无法通过消毒技术完全清除，从而对抗生素耐药性基因（ARG）的转移构成潜在风险。因此，为了更好地了解VBNC细菌并解释ARGs的潜在转移，本文系统地回顾了细菌进入VBNC状态后在形态、生理和毒力方面的变化。此外，本文还综述了VBNC细菌的定量检测方法，如细胞膜完整性介导的LIVE/DEAD Baclight检测法、基于qPCR的检测法和基于噬菌体的检测法，得出的结论是VBNC细菌仍缺乏原位和实时检测方法。然后评估了VBNC细菌的健康风险和环境应用价值，数据表明VBNC细菌在微生物利用领域具有巨大价值。此外，重点介绍了VBNC细菌的诱导条件（特别是通过光消毒）和形成机制。形成机制主要包括严格反应、一般应激反应系统和毒素-抗毒素（TA）系统。此外，评估了在基于光的消毒诱导的VBNC细菌形成期间和之后ARGs的水平基因转移（HGT）。研究发现ARGs可能通过结合、转化和转导进行转移。最后，总结了目前VBNC细菌转化的不足和未来的挑战，特别是受光消毒技术影响的不足和挑战。这篇综述为VBNC细菌的检测方法、形成机制、环境应用和潜在的ARG转移风险提供了新的见解。 
Abstract
Disinfection technologies, especially light-based disinfection, have undergone tremendous development and innovation, but this treatment can cause bacteria to enter viable but nonculturable (VBNC) state. Due to their strong tolerance, VBNC bacteria cannot be completely removed by disinfection technologies, thereby posing a potential risk for antibiotic resistance gene (ARG) transfer. Therefore, to better understand VBNC bacteria and interpret potential transfer of ARGs, this article systematically reviewed changes in morphology, physiology and virulence of bacteria after entering VBNC state. In addition, this article reviewed quantitative detection methods of VBNC bacteria, such as cell membrane integrity-mediated LIVE/DEAD Baclight assay, qPCR-based assays, and phage-based detection methods, concluding that there is still a lack of in-situ and real-time detection methods for VBNC bacteria. Health risks and environmental application value of VBNC bacteria were then valuated, with data indicating that VBNC bacteria have great value in the domain of microbial utilization. Furthermore, the induction conditions (especially by light-based disinfection) and formation mechanisms of VBNC bacteria were highlighted. Formation mechanisms mainly involve stringent response, general stress response system and toxin-antitoxin (TA) system. Moreover, horizontal gene transfer (HGT) of ARGs during and after the formation of VBNC bacteria induced by light-based disinfection was evaluated. It was found that ARGs may be transferred through conjugation, transformation and transduction. Finally, current deficiencies and future challenges for the transformation of VBNC bacteria, especially those influenced by light-based disinfection technologies, were summarized. This review provides new insights into detection methods, formation mechanisms, environmental applications and potential ARG transfer risks of VBNC bacteria.

https://www.tandfonline.com/doi/abs/10.1080/10643389.2021.1932397