Abstract
Bacteria navigate their way often as individual cells through their chemical and biological environment in aqueous medium or across solid surfaces. They swim when starved or in response to physical and chemical stimuli. Flagella-driven chemotaxis in bacteria has emerged as a paradigm for both signal transduction and cellular decision-making. By altering motility, bacteria swim toward nutrient-rich environments, movement modulated by their chemotaxis systems with the addition of pili for surface movement. The numbers and types of chemoreceptors reflect the bacterial niche and lifestyle, with those adapted to complex environments having diverse metabolic capabilities, encoding far more chemoreceptors in their genomes. The Alpha-proteobacteria typify the latter case, with soil bacteria such as rhizobia, endosymbionts of legume plants, where motility and chemotaxis are essential for competitive symbiosis initiation, among other processes. This review describes the current knowledge of motility and chemotaxis in six model soil bacteria: Sinorhizobium meliloti, Agrobacterium fabacearum, Rhizobium leguminosarum, Azorhizobium caulinodans, Azospirillum brasilense, and Bradyrhizobium diazoefficiens. Although motility and chemotaxis systems have a conserved core, rhizobia possess several modifications that optimize their movements in soil and root surface environments. The soil provides a unique challenge for microbial mobility, since water pathways through particles are not always continuous, especially in drier conditions. The effectiveness of symbiont inoculants in a field context relies on their mobility and dispersal through the soil, often assisted by water percolation or macroorganism movement or networks. Thus, this review summarizes the factors that make it essential to consider and test rhizobial motility and chemotaxis for any potential inoculant.
Highlights
There are several strategies bacteria use to actively navigate their environment, motile forces produced either through pili retraction or flagella rotation
Active bacterial motility tends to be controlled by chemotaxis systems that respond to different stimuli, allowing bacteria to migrate to optimal environments
A mutant of Azorhizobium caulinodans ORS571 with the chemotaxis cluster deleted was defective in Sesbania rostrata root surface colonization and competitive nodulation (Liu et al, 2018a)
Summary
There are several strategies bacteria use to actively navigate their environment, motile forces produced either through pili retraction or flagella rotation. The main chemotactic systems of S. meliloti, A. fabacearum, and R. leguminosarum have a similar core system to that of E. coli, belonging to class F7 and being associated with chemoreceptors of type 36H (Sourjik and Schmitt, 1998; Miller et al, 2007; Tambalo et al, 2010a,b; Wibberg et al, 2011) (see Figures 2A,B) These species do not encode CheZ, instead they encode two copies of CheY, with CheY2 propagating the signal and CheY1 acting as a phosphate sink, increasing the rate at which CheY2 returns to its unphosphorylated state (Sourjik and Schmitt, 1998). The che system is an F8-class cluster associated with chemoreceptors of type 34H and encodes an additional cheW gene with three chemoreceptors (Kaneko et al, 2002; Wuichet and Zhulin, 2010) The involvement of these systems on chemotaxis is currently unknown, it was found that only the subpolar flagellum of B. diazoefficiens responds chemotactically to glutamate and succinate (Quelas et al, 2016). Selecting for excess dispersal could be counterproductive by diluting inoculants away from the intended targets
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