Thursday, January 7, 2010

Double or nothing

As I finished my undergrad career and started thinking about graduate school, I was totally infatuated with the chromosomal speciation of treefrogs in the genus Hyla. Hyla versicolor and H. chrysoscelis, the 'gray treefrogs', have similar geographic distributions and look almost identical - except that one is a tetraploid version of the other. The increase in genome size is associated with a slight increase in cell size, which has trickle-down effects into physiology, the sound of their call, and other ecological factors, and of course they are reproductively isolated. As it turns out, Margaret Ptacek and colleagues were unraveling this mystery at the genetic level just as I was learning of it, and while I was disappointed not to be able to explore this for my graduate work, Margaret made up for it by paying for all the drinks when I visited Clemson a few years back.

So it was with considerable interest that I stumbled across one of the first tables of contents of the new year, in BMC Evolutionary Biology. Two co-occurring populations of the diatom Ditylum brightwellii, it turns out, differ in genome size. In this case, the belief is that there is a single taxonomic species harboring a very recent genome duplication polymorphism (which are likely cryptic species). Of course, a species by any other name... well, that's the problem isn't it? In the world of diatoms, according to Koester and colleagues, the 'barcode' standard is to use the 18S rDNA gene sequences and silica cell wall morphology in diagnosing species. However, already armed with evidence that two substantially distinct populations could be identified with the more rapidly-evolving ITS gene region, these researchers explored how differences in reproductive rates and size distributions might be associated with genome size.

See, diatoms are the petri dishes of the natural world. In order to reproduce, each side of the interlocking silica case separates and generates a new nested case. One of the offspring of this fission will be the same size as the parental individual, the other will be slightly smaller - the smaller of the two original cell walls, with an even smaller one nested within. At least that is how I understand it. Over time, these clonal lineages reduce substantially in size, and cell size is eventually limited by genome size; sexual reproduction then allows them to regain a larger cell size and the process repeats. So, the life history of this species requires an interesting interaction among the genome (which places a lower bound on cell size, and a lower bound on reproductive rate) and the population.

In Ditylum, Koester et al. were able to show that there are not only two very distinct genetic lineages, but that the one that is regionally localized to Puget Sound appears to have been generated through genome duplication. That is, there is a cosmopolitan species, and an offshoot lineage that was formed through some form of genome duplication, with concomitant changes in cell size, rates of population growth, and reproductive isolation. Koester et al. conclude that these lineages are cryptic species, and that this form of isolation may be common in marine diatoms.

More generally, this shows another way in which our understanding of biodiversity is changing rapidly thanks to molecular diversity analysis. The latest term to be coined by John Avise, biodiversity genetics, reflects the fact that we must now consider all of the new ways in which this technology can accelerate the rate of discovery in our natural world. Taxonomists trained in the morphology and phenotypic diversity of life are few; certainly too few to keep up with growing scientific collections, and the bottleneck in describing species can be a difficult one for management and conservation. The '18S or bust' approach in diatoms may be one standard that will change as more studies like this one, out of Armbrust's lab at Washington, illuminate how dynamic biological diversity can be.

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