[sci.chem] Molecular Phylogeny

chi9@quads.uchicago.edu (Lucius Chiaraviglio) (11/03/90)

In article <4578@husc6.harvard.edu> Ellington@Frodo.MGH.Harvard.EDU (Deaddog)
writes:
>In article <1990Oct30.030717.8923@midway.uchicago.edu> 
>chi9@quads.uchicago.edu (Lucius Chiaraviglio) writes:
>> combination of slow evolution and deep phylogenetic divergence,
>> which -- when the deeply-diverging organisms have a lot in common
>> phenotypically, which they do -- implies greater resemblence to ancestral
>> forms of life.
>   This is somewhat disturbing.  
>   First, isn't the fact that only a few, deep phylogenetic divergences 
>have been found between the *known* Archaebacteria a function of the 
>relatively small number that have been catalogued?  [. . .]

	While it is true that the number of Archaebacteria catalogued is
small, and that several Archaebacterial kingdoms are probably yet to be
discovered, a large number of Archaebacteria have been discovered.  Find a
review on Archaebacteria or some subgroup thereof which actually lists a bunch
of sample species (a somewhat out-of-date sample:  "Methanogens and the
Diversity of Archaebacteria" in a 1987 issue of _Microbiological Reviews_ --
sorry, I don't remember the authors' names or the exact issue off hand, but if
you want I could look it up when I get home).  Thus, while the picture is not
complete, it is definitely emerging.

	On the other hand, even though a much larger number of Eubacteria have
been characterized, some recent discoveries (Thermotoga for example) imply
that kingdoms remain to be discovered on this branch as well.

>   Second, while the Archaebacteria have some traits in common, their 
>phenotypic resemblances are immaterial to what a primitive life form looks 
>like; it is only  those traits that are held in common by each of the 
>three ur-kingdoms (or at least two out of three) that give us a logically 
>valid picture of the common ancestor of modern life (what I like to call 
>the progenote, but Woese has disagreed).

	That doesn't disagree with what I said (except that it seems better to
call the common ancestor a *protogenote* -- I think that's also what Woese
said).

>                                          Also, what makes you think that 
>Archaebacteria are more phenotypically similar within their ur-kingdom 
>than eukaryotes or eubacteria; what traits are you referring to?

	I didn't say this.  What I was getting at was that the features
(thermophily is the main one that I was thinking of) which the last common
ancestor of Archaebacteria has in common with the last common ancestor of
Eubacteria are more likely to be features of the last common ancestor of the
two lineages.  What I forgot to say in my original article was that since
recent molecular phylogeny (reference given below) indicates that Eukaryotes
diverge from the Archaebacteria after their common ancestor diverges from the
Eubacteria, this presumably applies to all three lineages.

>   Finally, the idea that the evolutionary rate of archaebacteria has been 
>slow

	Whoa -- I most certainly didn't say that.  I was talking about slower-
evolving and deep-diverging organisms WITHIN the Archaebacteria and the
Eubacteria, and did not say, or want to imply, that one group is as a whole
slower-evolving or has a sparser actual tree (as opposed to that which we have
catalogued) than the other.  (Actually, the Archaebacteria do fall into two
major subdivisions, which Woese calls the Crenarchaeota and Euryarchaeota,
whereas the Eubacteria have a more even branching pattern, and the Eukaryotes
form a subtree with strong apical dominance.)

>     seems unsupported:  aren't the ribosomal RNA sequences within the 
>archaebacterial lineage at least as divergent from each other as those 
>within the eubacterial or eukaryotic lineages?

	I didn't say they weren't.  However, the Eukaryotes really do seem to
be evolving at about twice the rate of the other two lineages -- see Carl
Woese's article in one of the June (+ or - 1 month) issues of _Proceedings of
the National Academy of Sciences_, and also the reference therein about how
the root of the overall tree of life was determined.

>                                                Doesn't the wide variety 
>of metabolic adaptations seen for different archaebacteria (halophiles, 
>thermophiles, acidophiles, chemolithotrophs, chemoautotrophs) demonstrate 
>that the rate of evolution has been at least as quick as in the other 
>lineages?

	No, it doesn't (although by molecular phylogenetic criteria the
Archaebacteria as a whole seem to be about as capable of rapid evolution as
the Eubacteria -- reference given above).  The Eubacteria have all of those
phenotypes -- EXCEPT METHANOGENESIS -- plus even more if you further subdivide
some of those that you listed above.  In contrast, the known Eukaryotes, which
have been evolving considerably more rapidly, have much less biochemical
diversity than the other two groups.

>   I may be misreading your intent, but it seems as though you want to 
>classify archaebacteria as 'living fossils'--and there just ain't such a 
>thing.
>A.E. 

	Everything has some living fossil aspect to it, but those organisms
which have evolved more slowly have more of it than others.  However, I was
NOT trying to classify the Archaebacteria as the prime living fossils.

--
|   Lucius Chiaraviglio    |    Internet:  chi9@midway.uchicago.edu