Effects of sublethal stress application on the survival of bacterial inoculants: a systematic review

The use of commercial bacterial inoculants formulated with plant-growth promoting bacteria (PGPB) in agriculture has shown significant prominence in recent years due to growth-promotion benefits provided to plants through different mechanisms. However, the survival and viability of bacterial cells in inoculants are affected during use and may decrease their effectiveness. Physiological adaptation strategies have attracted attention to solve the viability problem. This review aims to provide an overview of research on selecting sublethal stress strategies to increase the effectiveness of bacterial inoculants. The searches were performed in November 2021 using Web of Science, Scopus, PubMed, and Proquest databases. The keywords "nitrogen-fixing bacteria", "plant growth-promoting rhizobacteria", "azospirillum", "pseudomonas", "rhizobium", "stress pre-conditioning", "adaptation", "metabolic physiological adaptation", "cellular adaptation", "increasing survival", "protective agent" and "protective strategy" were used in the searches. A total of 2573 publications were found, and 34 studies were selected for a deeper study of the subject. Based on the studies analysis, gaps and potential applications related to sublethal stress were identified. The most used strategies included osmotic, thermal, oxidative, and nutritional stress, and the primary cell response mechanism to stress was the accumulation of osmolytes, phytohormones, and exopolysaccharides (EPS). Under sublethal stress, the inoculant survival showed positive increments after lyophilization, desiccation, and long-term storage processes. The effectiveness of inoculant-plants interaction also had positive increments after sublethal stress, improving plant development, disease control, and tolerance to environmental stresses compared to unappealed inoculants.