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Burkholderia glumae  causes bacterial panicle blight of rice and produces major virulence factors, including toxoflavin, under the control of the quorum-sensing (QS) system mediated by the  luxI  homolog,  tofI , and the  luxR  homolog,  tofR . In this study, a series of markerless deletion mutants of  B. glumae  for  tofI  and  tofR  were generated using the suicide vector system, pKKSacB, for comprehensive characterization of the QS system of this pathogen. Consistent with the previous studies by other research groups,  ΔtofI  and  ΔtofR  strains of  B. glumae  did not produce toxoflavin in Luria-Bertani (LB) broth. However, these mutants produced high levels of toxoflavin when grown in a highly dense bacterial inoculum (∼ 10 11  CFU/ml) on solid media, including LB agar and King’s B (KB) agar media. The  ΔtofI / ΔtofR  strain of  B. glumae , LSUPB201, also produced toxoflavin on LB agar medium. These results indicate the presence of previously unknown regulatory pathways for the production of toxoflavin that are independent of  tofI  and/or  tofR . Notably, the conserved open reading frame (locus tag: bglu_2g14480) located in the intergenic region between  tofI  and  tofR  was found to be essential for the production of toxoflavin by  tofI  and  tofR  mutants on solid media. This novel regulatory factor of  B. glumae  was named  tofM  after its homolog,  rsaM , which was recently identified as a novel negative regulatory gene for the QS system of another rice pathogenic bacterium,  Pseudomonas fuscovaginae . The  ΔtofM  strain of  B. glumae , LSUPB286, produced a less amount of toxoflavin and showed attenuated virulence when compared with its wild type parental strain, 336gr-1, suggesting that  tofM  plays a positive role in toxoflavin production and virulence. In addition, the observed growth defect of the  ΔtofI  strain, LSUPB145, was restored by 1 µM  N- octanoyl homoserine lactone (C8-HSL).

Dissection of Quorum-Sensing Genes in Burkholderia glumae Reveals Non-Canonical Regulation and the New Regulatory Gene tofM for Toxoflavin Production