Before the days of molecular phylogenies, the standard way of viewing microbial evolution was as process of physiological evolution: the ordering of the sequence of events in which different pathways that microbes use to harness carbon and energy arose. The physiological view of microbial evolution was, of course, replaced in the 1980s by a gene centered view of microbial evolution that was built around the ribosomal RNA tree of life, also called the universal tree or the three domain tree. The universal tree installed long sought order into microbial systematics, but left physiological evolution out in the cold, because physiology never mapped properly onto the rRNA tree. LGT decoupLes physioLoGy from phyLoGeny T he foregoing quote from Dickerson (1) typifies a physiological view of evolution, one in which lateral gene transfer between prokaryotes is normal and distributes metabolic capabilities among lineages. Almost four decades later, the quote is still quite modern. It embodies a particular approach to understanding microbial evolution, a view from the standpoint of physiology -the chemical reactions at the core of carbon and energy metabolism that drive the process of life forward. Physiology is important in evolution, whether carbon (2, 3), sulfur (4, 5), nitrogen (6), or the geochemical context of physiology (7). Dickerson's quote also puts a very natural and nonchalant 1980s mention of lateral gene transfer (LGT) into the picture of microbial evolution as a common component of genetic variation among prokaryotes, long before people were debating the significance of LGT in prokaryote evolution. It even goes so far as to say that
That was not because the universal tree had an incorrect branching pattern. Rather it was because physiological characters have never mapped neatly onto any phylogenetic tree for prokaryotes, regardless of its topology. The reason is that prokaryotes, though they have an undeniable tendency to vertically inherit their ribosome, distribute the physiological traits that enable synthesis of ribosomes via lateral gene transfer (LGT).LGT could possibly transform non-respiring lineages into respiring lineages via the transfer of many genes, something that we now know does actually occur during microbial evolution (8).Dickerson noted that the "evolutionary record" of microbes might have been scrambled. We have concepts about microbial evolution that are based on their evolutionary record, but what is the evolutionary record of microbes? What is it made of? It would appear that for microbial evolution there are only two kinds of records: geological and genomic. It is said that Earth records its own history (9), so do genomes. Putting geological and genomic evidence into a consistent picture of microbial evolution is a challenging undertaking, but that is the deliverable if we want a fuller picture of microbial evolution. Yet the only real connection between the two kinds of substance of the microbial evolutionary record -rocks and genes -is physiology. Physiology is arguably what...