Prokaryotes acquire genes from cells that are not their parents by three routes. Transformation: a cell takes up naked DNA from its surroundings (released by dead cells) and recombines it into its chromosome; some species are naturally competent, and it is the route by which pneumococci exchange capsule genes. Conjugation: a donor carrying a conjugative plasmid (the F factor of E. coli) builds a pilus, draws a recipient close, and passes a single strand of the plasmid through a pore while replicating it by the rolling circle; if the plasmid has integrated into the chromosome (an Hfr strain), chromosomal genes are transferred in order behind it, and the time at which each enters maps the chromosome. Transduction: a bacteriophage packages a piece of host DNA by mistake and injects it into the next cell it infects. Together these routes move resistance genes between species within hospitals in years, and have moved metabolic genes across the whole bacterial tree over geological time, so that a prokaryote’s ancestry is a web as much as a tree (Chapter 24).
Examples
Example 3.8 (Resistance on the move)
A resistance gene typically arises once, by mutation or from the soil bacterium that makes the antibiotic, and then travels: from a chromosome onto a transposon, from the transposon onto a conjugative plasmid, from the plasmid across species by conjugation and across strains by transduction, and back into a chromosome by transformation. Plasmids carrying five or six resistances at once (assembled in integrons, which capture gene cassettes) were found in Japan in the 1950s, a few years after the drugs came into use; the gene for the carbapenemase NDM-1, first seen in 2008, reached every continent within three years on a plasmid. The evolution of resistance is mostly not the evolution of new genes but the movement of old ones.