Structured habitats and the evolution of anticompetitor toxins in bacteria.

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Abstract
We demonstrate that in liquid cultures, defined in this study as a mass habitat, the outcome of competition between Escherichia coli that produce an antibacterial toxin (colicin) and sensitive E. coli is frequency dependent; the colicinogenic bacteria are at an advantage only when fairly common (frequencies in excess of 2 X 10(-2)). However, we also show that in a soft agar matrix, a structured habitat, the colicinogenic bacteria have an advantage even when initially rare (frequencies as low as 10(-6)). These contrasting outcomes are attributed to the colicinogenic bacteria's lower intrinsic growth rate relative to the sensitive bacteria and the different manner in which bacteria and resources are partitioned in the two types of habitats. Bacteria in a liquid culture exist as randomly distributed individuals and the killing of sensitive bacteria by the colicin augments the amount of resource available to the colicinogenic bacteria to an extent identical to that experienced by the surviving sensitive bacteria. On the other hand, the bacteria in a soft agar matrix exist as single-clone colonies. As the colicinogenic colonies release colicin, they kill neighboring sensitive bacteria and form an inhibition zone around themselves. By this action, they increase the concentration of resources around themselves and overcome their growth rate disadvantage. We suggest that structured habitats are more favorable for the evolution of colicinogenic bacteria.

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