Saturday, March 03, 2007

The Evolution of the HSP70 Gene Family

Here's a picture of my three second year project students in front of their poster. Blerta Kolaj, Milu Jauregui, and Zarna Shah collaborated on this project to study the evolution of various members of the HSP70 gene family.

Eukaryotes contain at at least five or six versions of HSP70 genes. Two of them are present in single copies (mostly) and they were present in the common ancestor of all eukaryotes. One of these is the mitochondrial version, which is derived from the proteobacterium ancestor of mitochondria. The other is a version found in the endoplasmic reticulum (ER). The ER member of the HSP70 gene family (=BiP) arose very early in eukaryotic evolution.

A phylogenetic tree of either the mitochondrial gene or the ER gene should reveal the evolution of all eukaryotic species. The two trees should be identical provided the genes evolve independently and there has been no horizontal gene transfer. Furthermore, the two trees will root each other since they join at a common ancestral node representing the split between eukaryotes and prokaryotes. The purpose of the experiment is to test this hypothesis and see if we have enough examples of ER and mitochondrial genes to make duplicate trees. The problem is that we need an example of each gene in every species we test. The fact that we now have quite a few genome sequences means that this has become more feasible than in the past.

Blerta, Milu, and Zarna were able to add in one more set of sequences; the chloroplast members of the HSP70 gene family. These genes are derived from the cyanobacteria ancestor of chloroplasts so they provide independent confirmation of the eukaryotic tree for plants and algae. Unfortunately, there aren't very many examples of species that have sequenced versions of the chloroplast gene, the mitochondrial gene, and the ER gene.

Here's the tree.

As you can see, the relationship of most species (actually genes) in the mitochondrial and ER branches is very similar. This confirms that the two family members are evolving independently. Differences occur with the protists such as trypanosomes, Leishmania, plasmodia, and cryptosporidium. This is unfortunate since analysis of HSP70 genes could have helped resolve the relationship of protists, which is very controversial. At least we're sure of the roots of each major branch—that's a big step forward.

The chloroplast and mitochondrial clades are closer together than either is to the eukaryotic ER versions of the HSP70 family. This is not unexpected since both organelle genes are prokaryotic in origin. It suggests that the universal root is along the branch leading to the ER clade.

Now my students have to turn their attention to their individual projects. They have about six weeks to finish up. I'll post summaries of their work in May. I'm pretty excited about all three projects, we could end up answering some pretty important questions using the HSP70 database.

7 comments:

  1. Very cool. I am curious, however, about what caused the very small discrepancies. I can think of several possibilities, (measurement accuracy or precision, different mutation rates in the 3 different genomes, etc), but I was wondering what the real (or at least the most probable) reason is in your opinion.

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  2. Hi, I am one of the students in the pic!
    I am not sure if we know the "real" reason, otherwise we would have tried to solve it right?
    We are somewhat dependent on the PHYLIP programs and we don't fully understand the algorithms that are used to produce the tree. also, Anything is possible when you are looking at something so backward in time, specially protists.
    But the closest explanation we have is, "long-branch attraction" hypothesis.
    Despite of all the possible discrepancies, we should also keep in mind that this could be "correct" position, but we don't know how it occured!

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  3. That would assume the real reason is solvable to begin with, which unfortunately isn't always the case. For instance precision-related errors due to the finite lengths of the genes in question may be fundamentally unavoidable (this is a common problem when dealing with any sort of discrete signal).

    And I wasn't meaning to imply that the results were wrong, just different (hence the use of the word "discrepancy" instead of "error"). As you said, they may really be different, or they may not. It would be interesting if they were different, since that might lead to new discoveries (of course being the same might as well).

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  4. zarna, nice job. but who is that weird guy sneaking in the background of your picture?

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  6. Great Article! It would be great to see the mitochondrial HSP70 tree overlaid with the ER HSP70 tree to track discrete deviations in pattern. Has there been any changes or additions to this tree since 2007? I would love to see an updated version, and perhaps (with your consent) post it to my HSP70 scientific resource guide http://www.hsp70.com.

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