The
three-domain system is a
biological classification introduced by
Carl Woese in 1990 that emphasizes his separation of
prokaryotes into two groups, originally called
Eubacteria and
Archaebacteria. Woese argued that, on the basis of differences in
16S rRNA genes, these two groups and the
eukaryotes each arose separately from an ancestral
progenote with poorly developed
genetic machinery. To reflect these primary lines of descent, he treated each as a
domain, divided into several different
kingdoms. The groups were also renamed the
Bacteria,
Archaea, and
Eukarya, further emphasizing the separate identity of the two prokaryote groups.
Although the three-domain system was quickly adopted by most
molecular systematists, biologists like
Ernst Mayr criticized him for over-emphasizing the uniqueness of the
archaebacteria and ignoring strong genetic similarities between the groups. Subsequent studies have confirmed that the archaea are unusual in the composition of their
cell membrane and structure of their
flagella. Other significant differences include archaeal systems for
DNA replication and
transcription which bear distinct similarity to those found in eukaryotes. For instance, archaeal
RNA polymerase consists of up to 14 subunits, whereas most bacterial RNA polymerases have only 4 subunits. Analysis of these subunits suggests that they're more closely related to those found in eukaryotes. Also, the archaea produce a number of
DNA-binding proteins with similarity to eukaryotic
histones.
Nevertheless, a minority viewpoint suggests retaining the older
two-empire system (Prokaryota and Eukaryota) and using the word
bacterium in its earlier meaning of
prokaryote.
Which system is preferable depends partly on the relationships of the
organisms in question. Although the
progenote hypothesis is discredited,
molecular trees tend to group living things into the three domains, with the eukaryotes placed beside or within the Archaea and the eubacteria forming a separate branch. However, it has been suggested this is an artifact of
long branch attraction and that the root may instead belong among the eubacteria, in which case many eubacterial lines diverged before the archaebacteria did.
In 2006, the discovery of unique properties of the
Mimivirus started a discussion about classification of (certain)
viruses as a fourth domain of life.
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