Volvox 2015: cell differentiation

One of the most studied aspects of Volvox development is the differentiation of its 2000 or so cells into two types: a few (usually 12-16) large reproductive cells (germ) and the rest small, biflagellate cells that provide motility (soma). The main genes controlling this differentiation have long been known, but the details of how they work are still being worked out.

Erik Hanschen (left) with Cristian Solari, David Smith, and Jillian Walker

Erik Hanschen (left) with Cristian Solari, David Smith, and Jillian Walker

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Volvox 2015: all about sex

I believe that sex is one of the most beautiful, natural, wholesome things that money can buy.

–Steve Martin

Volvox, and the volvocine algae in general, are well known as a model system for the evolution of multicellularity and cellular differentiation, but they are also an outstanding model for the evolution of sex and mating types. Volvocine algae are facultatively sexual, with haploid vegetative colonies reproducing asexually through mitosis but occasionally entering a sexual cycle that usually results in a diploid, desiccation-resistant zygote or ‘spore.’ Most of the small colonial species and unicellular relatives are isogamous, that is, the gametes are of equal size. Nevertheless, each species has two self-incompatible mating types, usually designated as ‘plus’ and ‘minus.’ In some of the larger species, the gametes have diverged into a small, motile form that we call sperm and a large, often immotile form that we call eggs. Across the eukaryotic domain, it is gamete size, not form of genitalia, fancy plumage, or receding hairline, that define males and females.

The volvocine algae span a wide range of mating systems, making them a useful (and I think underutilized) system for comparative studies of the evolution of sex. As I’ve already mentioned, both isogamous (equal-sized gametes) and oogamous (sperm and eggs) species exist, and there is good reason to suspect that oogamy has evolved independently in two separate lineages:

Isogamy and oogamy (Kirk, D.L. 2006. Oogamy: inventing the sexes. Curr. Biol., 16: R1028–R1030.)

Isogamy and oogamy (Kirk 2006. Curr. Biol., 16:R1028.)

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Volvox 2015

Today is day one of the Third International Volvox Conference in Cambridge, U.K. I arrived yesterday via Chicago, Ottowa, and a bus from Heathrow. Apparently there is a direct flight from Chicago to Heathrow; why I didn’t take that is a mystery. The accommodations at Wychfield are quite comfortable, although many of us have had the usual troubles figuring out English plumbing.

Last night was a pub dinner with half or so of the attendees; I had some very good fish & chips (good, but not as good as Go Fish).

Pub dinner with Stephanie Höhn, one of the organizers, in the foreground.

Pub dinner with Stephanie Höhn, one of the organizers, in the foreground.

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Fierce Roller will be offline next week

Missouri River Breaks

The Upper Missouri River National Monument in Montana. Photo by Bureau of Land Management.

I’ll be leaving Sunday morning for the Epic Canoe Trip on the Upper Missouri River (Coal Banks Landing to Kipp Recreation Area). We’ll be paddling, hiking, and camping in the White Cliffs of the Missouri and the Missouri River Breaks, a total of 107 miles. I’ll be totally offline; no cell service and no email. Don’t cry for me.

“…of the bignefs of a great corn of fand…”

Fig. 5 from van Leeuwenhoek 1700

“I thought convenient to get drawn one such before-mentioned particle, with the particles inclosed within it, as fig. 5 by E F sheweth. “

I usually try to comment on recent papers, but this time I’m going to go back a bit. More than a bit, really: 315 years, to what, as far as I know, is the first published report of Volvox (Van Leeuwenhoek, A. 1700. Part of a Letter from Mr Antony van Leeuwenhoek, concerning the Worms in Sheeps Livers, Gnats, and Animalcula in the Excrements of Frogs. Phil. Trans. Roy. Soc. London, 22:509–518). You know it’s old when half of the s’s look like f’s:

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(Probably not) Precambrian Volvox

A new(ish) paper in National Science Review evaluates the evidence for various interpretations of Ediacaran microfossils from the Weng’an biota in South China (Xiao et al. 2014. The Weng’an biota and the Ediacaran radiation of multicellular eukaryotes. Natl. Sci. Rev., 1:498–520.). I recommend checking it out; it’s open access, and there’s a lot of interesting stuff in there that I’m not going to address.

These fossils are undoubtedly multicellular, probably eukaryotic, and extremely enigmatic. Their age (582-600 million years) means they could have important implications for the evolution of multicellularity, and their exceptional preservation in great numbers creates the potential for reconstructing their life cycles in great detail. Some of the Weng’an fossils have been interpreted as volvocine algae, an interpretation that I find highly unlikely.

Some of the Weng’an fossils are thought to represent red algae, and this would not be terribly surprising, since red algae have been around for at least 1.2 billion years. Others, for example the tubular fossils, are more problematic, with interpretations as diverse as cyanobacteria, eukaryotic algae, crinoids, and cnidarians.

Fig. 8 from Xiao et al. 2014

Figure 8 from Xiao et al. 2014: Schematic diagram showing diagnostic features of the five recognized species of tubular microfossils in the Weng’an biota.

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Evolution of eusociality

Last month, two papers on the evolution of eusociality were published in high-profile journals: one by Karen M. Kapheim and colleagues in Science, the other by Sandra M. Rehan and Amy L. Toth in Trends in Ecology & Evolution (TREE). Social and eusocial insects are an attractive system for studying major transitions, sharing some of the key features that make the volvocine algae so good for this purpose: multiple, independent origins of traits thought to be important to the transition and extant species with intermediate levels of sociality. These features make the social insects, like the volvocine algae, well-suited for comparative studies.
Figure 1 from Rehan & Toth: (A) Overview of phylogeny of aculeate Hymenoptera (with the nonhymenopteran but eusocial termites as an outgroup), highlighting independent origins of sociality (colored branches), groups with species ranging from solitary to primitively social (green), primitively social to advanced eusocial (orange), solitary to advanced eusocial (blue), and all species advanced eusocial (grey). (B) The full range of the solitary to eusocial spectrum (blue) and predictions of which genomic mechanisms are hypothesized to operate at different transitional stages of social evolution (broken arrows).

Figure 1 from Rehan & Toth: (A) Overview of phylogeny of aculeate Hymenoptera (with the nonhymenopteran but eusocial termites as an outgroup), highlighting independent origins of sociality (colored branches), groups with species ranging from solitary to primitively social (green), primitively social to advanced eusocial (orange), solitary to advanced eusocial (blue), and all species advanced eusocial (grey). (B) The full range of the solitary to eusocial spectrum (blue) and predictions of which genomic mechanisms are hypothesized to operate at different transitional stages of social evolution (broken arrows).

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An Ode to Unicellularity

Biosphere 2

Biosphere 2, the site of the First International Volvox Meeting in 2011.

This year’s Volvox meeting, as with the previous two, will feature an image/video/arts competition. Erik Hanschen, a graduate student in the Michod lab, has kindly granted me permission to post the winning entry in the poetry contest at the first Volvox meeting: a sonnet in honor of Chlamydomonas.

An Ode to Unicellularity – Erik Hanschen

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