Frederick Cohan—the Origins of Ecological Diversity in Prokaryotes

How do we distinguish bacterial species? Cohan shows us some nice diagrams of phenotypic and molecular clusters, and they show groups separated by gaps — therefore, species. Unfortunately the species defined thereby are big and contain considerable diversity within them. Darwin defined species as divergent forms between which one finds morphological gaps. Mayr: cohesive set of organisms whose divergence is constrained by genetic exchange. Speciation requires a breakdown of that exchange.

Mallet has developed a version of Darwin's species definition that includes molecular characters. Under Mayr, speciation is tough, under Mallet, speciation is easy. The two models differ in the frequency of cladogenesis.

How do bacterial species maintain cohesion? Periodic selection purges divergent populations. Diversity within ecotypes is maintained by selective purges, but ecotypes that found new populations in new environments will not face the same selective effects.

Why doesn't the free(er) exchange of genetic material between bacterial species lead to a convergence or fusion of species? One reason is the rarity of genetic exchange. If two ecotypes have a suite of niche-specifying genes, low frequency of interchange will not be sufficient to prevent divergence.This does not prevent free exchange of niche-transcending genes, genes that are useful in different environments.

Lots of details from Cohan's work followed, and I confess to being a bit lost in places. He's looking at different soil bacteria that are found in different ecotypes—for instance, having different characteristic fatty acid content depending on whether they are found on a north-facing or south-facing slope. He argues that speciation is easy because genetic exchange doesn't prevent speciation. They're working on finding and confirming ecotypes with whole genome sequencing.

There is cohesion with local populations in one niche, but there is also niche-specifying divergence that is in defiance of cohesion. In animals and plants, niche-transcending genes are only shared between close relatives; in bacteria, they can be shared by the most distant relatives. This sharing doesn't interfere with divergence.