in article [email hidden], Jim Menegay at
[email hidden] wrote on 4/16/04 9:29 PM:
Quoted message said:Tim Tyler <[email hidden]> wrote in message
news:<[email hidden]>...
Quoted message said:Robert Karl Stonjek <[email hidden]> wrote or
quoted:
Quoted message said:There has been a lot of talk about comparisons between
the chimp and human genome, but what work is being done
on human-bonobo genomes?
It seems to me that we are much closer to the Bonobo -
they have smaller noses than chimps; pink lips; hairless
faces; can walk upright carrying items in their hands
including food; are less aggressive than Chimps; have
Quoted message said:Quoted message said:Quoted message said:appetite to humans.
Does anyone have more info on this?
We branched from them at the same time as we branched
from chimps.
However, it is widely speculated that the LCA was more
like a bonobo than a chimpanzee.
These ideas raise a number of questions in my mind, not so
much related to chimps, bonobos, and humans per se, but
related to theories of drift, speciation, and genomics.
The best way to ask these questions seems to me to present
a series of fictitious comments regarding Stonjek's ideas
that might arise from non-experts in sbe, and then to ask
for a critique of these comments by the experts.
I'll take the bait.
Here goes:
Quoted message said:
1. I can't believe that all those similarities between
Bonobo and [censored] could arise by convergence (ie. Bonobo
following in [censored]'s footsteps after a several million
year delay.) Therefore, I have to assume that many of
these traits were also present in the LCA of [censored] and
Bonobo.
OK, but you don't have to assume your way out of an
uncomfortable hypothesis. You could simply posit a more
comforting one, like you essentially did at the end. Of
course, there is no guarantee that the data will end up
supporting your comfort level, but then again that is how
learning takes place.
Quoted message said:2. You also have to assume that they were present in the
LCA of Bonobo and Chimp, but they were then lost on
the Chimp branch.
This would indeed be an extension of the hypothesis
outlined above.
Quoted message said:3. But that means that, since the divergence of Chimp and
Bonobo, Chimp has been evolving faster.
It would indeed mean this.
Quoted message said:4. A lot of those differences between Chimp and Bonobo
don't look like adaptations. So, I'm guessing that
this means that they arose by drift.
This would be an independent hypothesis. The validity of the
previous hypothesis would not depend upon the validity of
this "guess."
Quoted message said:5. The hypothesis that Chimp is drifting faster should be
checkable just by comparing the Chimp and Bonobo
genomes.
There are a number of things you could compare directly
between these gene pools as tests of this hypothesis. There
are a number of freeware programs available, for example,
that examine single gene pools (not in comparison with other
gene pools) for the "fingerprints" of drift, such as the
ratio of microsatellite size ranges to the number of alleles
with different sizes at a particular locus. Although I am
not aware of methods designed specifically for the kind of
comparison suggested here, you could easily compare test
statistics generated from the single gene pool tests.
Quoted message said:6. No, to make that test, you need an outgroup. You need
to include the [censored] genome in this analysis.
Although you do not NEED to do this, phylogenetically
grounded methods of comparison provide a potentially
powerful approach. The problem with power is that it can be
misused (misleading), so you have to know what you are doing
here. Outgroups are generally used as surrogates for the
true MRCA of the ingroup (chimps and bonobos, in this case).
If the outgroup is different from the ingroup MRCA in key
aspects, then you can be misled by the analysis. This issue
is not as big of a problem with many of the relative rate
tests, although it is still something to be considered.
Quoted message said:7. In any case, if Chimp has drifted faster since the
divergence from Bonobo, then there has to be an
explanation. I think it could be due to a series of
bottlenecks in the Chimp population. Low population
increases drift.
Right. Genetic diversity is lost from a population faster by
drift when the effective population size is lower.
Quoted message said:8. No. That is wrong. Drift speed is independent of
population. It would have to be because Chimp has a
higher mutation rate than Bonobo.
7 is not wrong, but neither is 8; one focuses on short term
effects, while the other focuses on long term effects. When
drift acts fast enough, or when the strength of selection is
low enough, then the evolution of the population is more
strongly controlled by the mutation rate. As Kimura pointed
out, in the mutation/drift balance model population size has
no effect on the rate of evolutionary substitution (allelic
fixation) in a population. Smaller populations drift faster,
but have to wait longer for new mutations to generate
polymorphism. Larger populations have plenty of new
mutations to deal with every generation, but drift happens
more slowly. These effects negate the influence of
population size on the rate of evolutionary substitution.
Quoted message said:9. Yes, but is it higher than the mutation rate in the
Chimp-Bonobo LCA? I don't think it can be. Mutation
rates cannot increase in evolution. Therefore we would
have to assume that the Bonobo somehow decreased its
mutation rate.
I have never seen an argument positing that mutation rates
cannot increase in evolution, and this claim seems patently
false to me.
Quoted message said:10. But can't we tell whether it was an increase in Chimp
mutations or a decrease in Bonobo mutations by looking
at the genome data?
I don't think this would be possible by looking directly at
these genomes. You would probably need to use a
phylogenetically grounded comparative method to get at this.
Quoted message said:11. Maybe, but we would need another outgroup - Gorilla,
or better yet, Pongo.
Yup. In fact, I think you would need to nest the
chimp/bononbo clade into a fairly rich phylogenetic context
to characterize the history of mutation rate evolution in
the lineage underpinning the chimp/bonobo MRCA.
Quoted message said:12. You molecular people are giving me a headache. Isn't
it obvious that the traits that Stonjek cites as
differences between Chimp and Bonobo are mostly the
kinds of traits that might be involved in
Quoted message said:Bonobo by a kind of species budding. And that Chimp
quickly developed differences in those traits to enforce
reproductive isolation.
This hypothesis could be roughly true, even if the other
hypotheses are roughly true. It approaches the issue from a
different angle.
Quoted message said:13. That is OK for all of the traits except walking
upright and carrying things in your hand. It is crazy
to claim that that trait already existed in the Chimp-Human-
Bonobo LCA, isn't it? I mean, don't we have skeletons
refuting this? So, that trait, at least, arose by
convergence.
I see this argument as sloppy on many points, and
therefore unconvincing to a skeptic. Note that I am not
arguing it is false, just that it is based on too much arm
waiving. It would not be "crazy to claim that that trait
already existed in the Chimp-Human-Bonobo LCA;" this would
just be another hypothesis. It seems unlikely to be true,
but that would make a bold hypothesis, which is a good
thing. We do not have skeletons refuting this hypothesis
at this point. In fact, we have very little homonid fossil
evidence at all from the critical period of time when the
lineages leading to humans, gorillas, chimps and bonobos
became isolated. There may be a few teeth from that time
period, but I don't think there is ANY other fossil
material of value to point to.
Cheers,
Guy Hoelzer