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After observing these patterns
of diversity and its distribution and their correspondence to ideas
about plant and animal speciation, I took a sabbatical leave in 2000 to
the lab of Fred Cohan’s [http://www.wesleyan.edu/bio/cohan/cohan.html]
to learn about evolutionary theory. The empirical results we observed
in our studies match well with periodic selection theory
[Cohan,
2002] (Fig. 9). The idea is that populations, which have
variation, evolve through a succession of periodic sweeps of diversity.
One most-fit variant out-competing all others and carrying forward the
genetic know how to occupy the population’s niche. However, whenever a
variant arises which has a novel ecology (occupies a different niche
than the parental population) it is no longer selected in the same way
(i.e., survives periodic sweeps of the parental population). It is then
free to diverge from the parental population, giving rise to a new
population which undergoes its own private periodic sweeps. The
eventual result is two ecologically distinct populations. Geographic
isolation can have the same effect as ecological adaptation in driving
populations apart. Fred has shown that a high-resolution molecular
technique for analyzing population genetics has potential to detect these terminal ecotypical clusters. The method, called
Multi-Locus Sequence Typing (MLST),
involves PCR amplification of 7 rapidly evolving protein encoding genes
and sorting variants into clonal complexes. Fred has developed an
evolutionary simulation that suggests that MLST clonal complexes equate
to putative ecotypes. In a project sponsored by the NSF
Frontiers in
Integrative Biological Research (FIBR)
and NASA Exobiology Programs, Fred and I plan to develop a
cultivation-independent MLST approach to study, at very high resolution,
Synechococcus ecotypes within the mat. As a part of the FIBR project,
we will collaborate with
John Heidelberg of the Institute for Genomic
Research (TIGR)
to obtain genomic sequences of two Synechococcus isolates that are
genetically relevant to the mats we study. John will also conduct
direct environmental genomic analysis of predominant mat populations, as
a theory-independent means of investigating how genomic diversity may be
organized into populations. Genomic sequence data will permit us, in
collaboration with
Devaki Bhaya
and
Arthur Grossman
(Carnegie Institution/Stanford), to develop microarrays that will be
used to investigate gene expression in situ within the mats.
Ultimately, we hope to compare the distributions of allelic variants of
highly expressed genes, with alleles that mark MLST clonal complexes
(putative ecotypes). |
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Figure 9. Periodic Selection
Theory. |
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