The regulated genes with putative function Among the 302 genes significantly altered in transcription by root exudates, 44 were annotated to encode a putative enzyme or a hypothetical protein. Similar to the genes with known function, these 44 genes fell into three categories: metabolism of carbohydrates and related molecules, metabolism of amino acids and related PLX4032 chemical structure molecules, and transport/binding proteins and lipoproteins (Additional file 1: Table S2). Some of the 44 genes were closely associated with plant-microbe interactions. For example, the transcription of ydjL, nowadays
renamed bdhA, encoding acetoin reductase/butanediol dehydrogenase [53], was 1.5-fold enhanced by root exudates. 2, 3-Butanediol is a volatile organic compound released by PGPR and able to promote significantly plant growth [54]. The expression of the gene AZD1390 epsE, residing in a 15-gene operon epsA-O, was also enhanced by root exudates. EpsE is involved in formation of biofilm by arresting flagellar rotation of cells embedded in biofilm matrix [55]. Another activated gene was dfnY, which encodes a hypothetical protein. Like other induced genes known to be involved in antibiotic production such as dfnF dfnG dfnI and dfnJ (Table 3), dfnY is part of the gene cluster responsible for synthesis
of the polyketide antibiotic difficidin. It is worth mentioning that antibiotic production is energetically very costly and its strict control is a clear evolutionary advantage. In contrast
to a few genes significantly altered during the exponential phase (OD1.0), hundreds of genes were differentially expressed in presence of root exudates during transition to stationary growth phase (OD3.0). Such a difference may not be surprising. The transcription of most bacterial genes during the exponential growth phase is typically initiated by RNA polymerase holoenzyme carrying the housekeeping transcription factor σA, while in the stationary phase, transcription is mainly accomplished by RNAP carrying alternative sigma factors allowing to adapt to a permanently changing environment. The extracytoplasmic-function (ECF) sigma factor W was enhanced in presence of root-exudate (Figure 5). SigW is known as being expressed Thymidylate synthase in early stationary growth-phase and induced by various cell wall antibiotics, alkaline shock, and other stresses affecting the cell envelope. It controls a large “antibiosis” regulon involved in mediating resistance to various antibiotics including fosfomycin and the antibiotic Trichostatin A peptides sublancin and SdpC [56]. It has been observed that many virulence-associated factors influence the colonization, persistence and spreading mechanisms of the human pathogen Streptococcus pyogenes in a growth phase-dependent manner [57–59]. Likewise, rhizobacteria may employ an early stationary phase-related mechanism to favor expression of those genes that mediate rhizosphere competence.
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