Quorum sensing (QS) bacteria regulate gene expression
collectively
by exchanging diffusible signal molecules known as autoinducers. Although
QS is often studied in well-stirred laboratory cultures, QS bacteria
colonize many physically and chemically heterogeneous environments
where signal molecules are transported primarily by diffusion. This
raises questions of the effective distance range of QS and the degree
to which colony behavior can be synchronized over such distances.
We have combined experiments and modeling to investigate the spatiotemporal
patterns of gene expression that develop in response to a diffusing
autoinducer signal. We embedded a QS strain in a narrow agar lane
and introduced exogenous autoinducer at one terminus of the lane.
We then measured the expression of a QS reporter as a function of
space and time as the autoinducer diffused along the lane. The diffusing
signal readily activates the reporter over distances of ∼1
cm on time scales of ∼10 h. However, the patterns of activation
are qualitatively unlike the familiar spreading patterns of simple
diffusion, as the kinetics of response are surprisingly insensitive
to the distance the signal has traveled. We were able to reproduce
these patterns with a mathematical model that combines simple diffusion
of the signal with logistic growth of the bacteria and cooperative
activation of the reporter. In a wild-type QS strain, we also observed
the propagation of a unique spatiotemporal excitation. Our results
show that a chemical signal transported only by diffusion can be remarkably
effective in synchronizing gene expression over macroscopic distances.
Sinorhizobium meliloti growing on soft agar can exhibit an unusual surface spreading behaviour that differs from other bacterial surface motilities. Bacteria in the colony secrete an exopolysaccharide-rich mucoid fluid that expands outward on the surface, carrying within it a suspension of actively dividing cells. The moving slime disperses the cells in complex and dynamic patterns indicative of simultaneous bacterial growth, swimming and aggregation. We find that while flagellar swimming is required to maintain the cells in suspension, the spreading and the associated pattern formation are primarily driven by the secreted exopolysaccharide EPS II, which creates two entropy-increasing effects: an osmotic flow of water from the agar to the mucoid fluid and a crowding or depletion attraction between the cells. Activation of these physical/chemical phenomena may be a useful function for the high molecular weight EPS II, a galactoglucan whose biosynthesis is tightly regulated by the ExpR/SinI/SinR quorum-sensing system: unlike bacterial colonies that spread via bacterium-generated, physical propulsive forces, S. meliloti under quorum conditions may use EPS II to activate purely entropic forces within its environment, so that it can disperse by passively 'surfing' on those forces.
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