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Summary: Rhizomicrobiome plays important roles in plant growth and health, contributing to the sustainable development of agriculture. Plants recruit and assemble the rhizomicrobiome to satisfy their functional requirements, which is widely recognized as the 'cry for help' theory, but the intrinsic mechanisms are still limited.In this study, we revealed a novel mechanism by which plants reprogram the functional expression of inhabited rhizobacteria, in addition to the de novo recruitment of soil microbes, to satisfy different functional requirements as plants grow. This might be an efficient and low‐cost strategy and a substantial extension to the rhizomicrobiome recruitment theory.We found that the plant regulated the sequential expression of genes related to biocontrol and plant growth promotion in two well‐studied rhizobacteria Bacillus velezensis SQR9 and Pseudomonas protegens CHA0 through root exudate succession across the plant developmental stages. Sixteen key chemicals in root exudates were identified to significantly regulate the rhizobacterial functional gene expression by high‐throughput qPCR.This study not only deepens our understanding of the interaction between the plant–rhizosphere microbiome, but also provides a novel strategy to regulate and balance the different functional expression of the rhizomicrobiome to improve plant health and growth. [ABSTRACT FROM AUTHOR]

Efficient root colonization of plant growth‐promoting rhizobacteria is critical for their plant‐beneficial functions. However, the strategy to overcome plant immunity during root colonization is not well understood. In particular, how Bacillus strains cope with plant‐derived reactive oxygen species (ROS), which function as the first barrier of plant defence, is not clear. In the present study, we found that the homolog of flg22 in Bacillus velezensis SQR9 (flg22SQR9) has 78.95% identity to the typical flg22 (flg22P.s.) and induces a significant oxidative burst in cucumber and Arabidopsis. In contrast to pathogenic or beneficial Pseudomonas, live B. velezensis SQR9 also induced an oxidative burst in cucumber. We further found that B. velezensis SQR9 tolerated higher H2O2 levels than Pst DC3000, the pathogen that harbours the typical flg22, and that it possesses the ability to suppress the flg22‐induced oxidative burst, indicating that B. velezensis SQR9 may exploit a more efficient ROS tolerance system than DC3000. Further experimentation with mutagenesis of bacteria and Arabidopsis showed that the two‐component regulatory system, ResDE, in B. velezensis SQR9 is involved in tolerance to plant‐derived oxidative stress, thus contributing to root colonization. This study supports a further investigation of the interaction between beneficial rhizobacteria and plant immunity. The beneficial rhizobacterium Bacillus velezensis SQR9 or its flg22 peptides triggered oxidative burst in cucumber and Arabidopsis. The two‐component regulatory system, ResDE, in B. velezensis SQR9 is involved in tolerance to plant‐derived oxidative stress, thus contributing to root colonization. [ABSTRACT FROM AUTHOR]

The phytohormone indole‐3‐acetic acid (IAA) has been demonstrated to contribute to the plant growth‐promoting effect of rhizobacteria, but the IAA biosynthesis pathway in rhizobacteria remains unclear. The ysnE gene, encoding a putative tryptophan acetyltransferase, has been demonstrated to be involved in and strongly contribute to IAA production in Bacillus, but the mechanism is unknown. In this study, to investigate how ysnE participates in IAA biosynthesis in the plant growth‐promoting rhizobacterium Bacillus amyloliquefaciens SQR9, differences in the produced IAA biosynthesis intermediates between wild‐type SQR9 and ΔysnE were analyzed and compared, and the effects of different intermediate compounds on the production of IAA and the accumulation of other intermediates were also investigated. The results showed that the mutant ΔysnE produced more indole‐3‐lactic acid (ILA) and tryptamine (TAM) than the SQR9 wild‐type strain (nearly 1.6‐ and 2.1‐fold), while the production of tryptophol (TOL) was significantly decreased by 46%. When indole‐3‐pyruvic acid (IPA) served as the substrate, the concentration of ILA in the ΔysnE fermentation broth was much higher than that of the wild type, while IAA and TOL were significantly lower, and ΔysnE was lower than SQR9 in IAA and TOL with the addition of TAM. The TOL content in the ΔysnE fermentation broth was much lower than that in the wild‐type SQR9 with the addition of ILA. We suggest that ysnE may be involved in the IPA and TAM pathways and play roles in indole acetaldehyde (IAAld) synthesis from IPA and TAM and in the conversion of ILA to TOL. [ABSTRACT FROM AUTHOR]

Lateral root formation is coordinated by both endogenous and external factors. As biotic factors, plant growth‐promoting rhizobacteria can affect lateral root formation, while the regulation mechanism is unclear. In this study, by applying various marker lines, we found that volatile compounds (VCs) from Bacillus amyloliquefaciens SQR9 induced higher frequency of DR5 oscillation and prebranch site formation, accelerated the development and emergence of the lateral root primordia and thus promoted lateral root development in Arabidopsis. We demonstrated a critical role of auxin on B. amyloliquefaciens VCs‐induced lateral root formation via respective mutants and pharmacological experiments. Our results showed that auxin biosynthesis, polar transport and signalling pathway are involved in B. amyloliquefaciens VCs‐induced lateral roots formation. We further showed that acetoin, a major component of B. amyloliquefaciens VCs, is less active in promoting root development compared to VC blends from B. amyloliquefaciens, indicating the presence of yet uncharacterized/unknown VCs might contribute to B. amyloliquefaciens effect on lateral root formation. In summary, our study revealed an auxin‐dependent mechanism of B. amyloliquefaciens VCs in regulating lateral root branching in a non‐contact manner, and further efforts will explore useful VCs to promote plant root development. The volatile compounds from Bacillus spp. increase the frequency of prebranch site formation and further accelerated the emergence of lateral root primordia (LRP) from the primary root (PR) in Arabidopsis, resulting in a denser lateral root (LR) along the PR spatially. This process is dependent on auxin signaling pathway; fully functional machinery of YUCs‐mediated auxin biosynthesis and polar auxin transport are also required. [ABSTRACT FROM AUTHOR]

Chemotaxis towards root exudates and subsequent biofilm formation are very important for root colonization and for providing the beneficial functions of plant growth-promoting rhizobacteria (PGPRs). In this study, in comparison with other root-secreted compounds, d-galactose in the root exudates of cucumber was found to be a strong chemoattractant at the concentration of 1 μM for Bacillus velezensis SQR9. Chemotaxis assays with methyl-accepting chemotaxis proteins (MCPs) deletion strains demonstrated that McpA was solely responsible for chemotaxis towards d-galactose. Interestingly, d-galactose significantly enhanced the biofilm formation of SQR9 in an McpA-dependent manner. Further experiment showed that d-galactose also enhanced root colonization by SQR9. In addition, the secretion of d-galactose by cucumber roots could be induced by inoculation with SQR9, indicating that d-galactose may be an important signal in the interaction between plant and SQR9. These findings suggested that the root-secreted d-galactose was a signal, the secretion of which was induced by the beneficial bacteria, and which in turn induced colonization of the bacteria. [ABSTRACT FROM AUTHOR]

Aside from applications in the production of commercial enzymes and metabolites, Bacillus amyloliquefaciens is also an important group of plant growth-promoting rhizobacteria that supports plant growth and suppresses phytopathogens. A host-genotype-independent counter-selectable marker would enable rapid genetic manipulation and metabolic engineering, accelerating the study of B. amyloliquefaciens and its development as both a microbial cell factory and plant growth-promoting rhizobacteria. Here, a host-genotype-independent counter-selectable marker pheS was constructed through a point mutation of the gene pheS, which encodes the α-subunit of phenylalanyl-tRNA synthetase in Bacillus subtilis strain 168. In the presence of 5 mM p-chloro-phenylalanine, 100 % of B. amyloliquefaciens strain SQR9 cells carrying pheS were killed, whereas the wild-type strain SQR9 showed resistance to p-chloro-phenylalanine. A simple pheS and overlap-PCR-based strategy was developed to create the marker-free deletion of the amyE gene as well as a 37-kb bmy cluster in B. amyloliquefaciens SQR9. The effectiveness of pheS as a counter-selectable marker in B. amyloliquefaciens was further confirmed through the deletion of amyE genes in strains B. amyloliquefaciens FZB42 and NJN-6. In addition, the potential use of pheS in other Bacillus species was preliminarily assessed. The expression of PheS in B. subtilis strain 168 and B. cereus strain ATCC 14579 caused pronounced sensitivity of both hosts to p-chloro-phenylalanine, indicating that pheS could be used as a counter-selectable marker (CSM) in these strains. [ABSTRACT FROM AUTHOR]

Chemotaxis, the ability of motile bacteria to direct their movement in gradients of attractants and repellents, plays an important role during the rhizosphere colonization by rhizobacteria. The rhizosphere is a unique niche for plant–microbe interactions. Root exudates are highly complex mixtures of chemoeffectors composed of hundreds of different compounds. Chemotaxis towards root exudates initiates rhizobacteria recruitment and the establishment of bacteria–root interactions. Over the last years, important progress has been made in the identification of root exudate components that play key roles in the colonization process, as well as in the identification of the cognate chemoreceptors. In the first part of this review, we summarized the roles of representative chemoeffectors that induce chemotaxis in typical rhizobacteria and discussed the structure and function of rhizobacterial chemoreceptors. In the second part we reviewed findings on how rhizobacterial chemotaxis and other root–microbe interactions promote the establishment of beneficial rhizobacteria-plant interactions leading to plant growth promotion and protection of plant health. In the last part we identified the existing gaps in the knowledge and discussed future research efforts that are necessary to close them. [ABSTRACT FROM AUTHOR]