Spread and transmission of bacterial pathogens in experimental populations of the nematode Caenorhabditis elegans.

Caenorhabditis elegans is frequently used as a model species for the study of bacterial virulence and innate immunity. In recent years, diverse mechanisms contributing to the nematode's immune response to bacterial infection have been discovered. Yet despite growing interest in the biochemical and molecular basis of nematode-bacterium associations, many questions remain about their ecology. Although recent studies have demonstrated that free-living nematodes could act as vectors of opportunistic pathogens in soil, the extent to which worms may contribute to the persistence and spread of these bacteria has not been quantified. We conducted a series of experiments to test whether colonization of and transmission between C. elegans nematodes could enable two opportunistic pathogens (Salmonella enterica and Pseudomonas aeruginosa) to spread on agar plates occupied by Escherichia coli. We monitored the transmission of S. enterica and P. aeruginosa from single infected nematodes to their progeny and measured bacterial loads both within worms and on the plates. In particular, we analyzed three factors affecting the dynamics of bacteria: (i) initial source of the bacteria, (ii) bacterial species, and (iii) feeding behavior of the host. Results demonstrate that worms increased the spread of bacteria through shedding and transmission. Furthermore, we found that despite P. aeruginosa's relatively high transmission rate among worms, its pathogenic effects reduced the overall number of worms colonized. This study opens new avenues to understand the role of nematodes in the epidemiology and evolution of pathogenic bacteria in the environment.

Some C. elegans and bacteria strains were provided by the Caenorhabditis Genetics Centre, which is funded by NIH Office of Research Infrastructure Programs (P40 OD010440). We thank Andrew Grant and Craig Winstanley for providing strains and reagents. We thank Mark Viney and two anonymous reviewers for helpful comments on the manuscript. This research was funded by a grant from the Biotechnology and Biological Sciences Research Council (grant number BB/I012222/1) to OR. OR also acknowledges funding from the Royal Society (University Research Fellowship).

This is the author accepted manuscript. The final published version can be found on the publisher's website at: http://aem.asm.org/content/early/2014/06/23/AEM.01037-14.long Copyright © 2014, American Society for Microbiology. All Rights Reserved