New review of green algal sex

Hiroyuki Sekimoto from Japan Women’s University has published a review of sexual reproduction in the volvocine algae and in the Charophyte Closterium in the Journal of Plant Research. In addition to a brief description of the Chlamydomonas sexual cycle, it includes a succinct review of the genetics of sex and sex determination. Unfortunately, the article is paywalled, and my inquiry to the author has so far gone unanswered.

Figure 1 from Sekimoto 2017. The life cycle of Chlamydomonas reinhardtii. Vegetative cells (V) di erentiate into mt+ and mt− gametes (G) during nitrogen starvation (−N). Mating types are restricted by mating-type loci (+ and −). When gametes are mixed, the plus and minus agglutinin mol- ecules on their agellar surfaces adhere to each other, and this adhe- sion results in increased intracellular cAMP levels. The signal trig- gers gamete cell wall release and mating-structure activation. Cells then fuse to form binucleate quadri agellated cells. Zygotes with thick cell walls germinate in response to light and nitrogen supple- mentation, and undergo meiosis to release four haploid vegetative cells

Figure 1 from Sekimoto 2017. The life cycle of Chlamydomonas reinhardtii. Vegetative cells (V) differentiate into mt+ and mt− gametes (G) during nitrogen starvation (−N). Mating types are restricted by mating-type loci (+ and −). When gametes are mixed, the plus and minus agglutinin molecules on their flagellar surfaces adhere to each other, and this adhesion results in increased intracellular cAMP levels. The signal triggers gamete cell wall release and mating-structure activation. Cells then fuse to form binucleate quadriflagellated cells. Zygotes with thick cell walls germinate in response to light and nitrogen supplementation, and undergo meiosis to release four haploid vegetative cells.

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J. S. Huxley part 2: Volvox

Last time, I wrote about Julian Huxley’s 1912 book, The Individual in the Animal Kingdom, and his use of the volvocine algae as an example. I liked most of what he had to say, though I took issue with his assertion that

…all the other members of the family except Volvox…are colonies and nothing more—their members have united together because of certain benefits resulting from mere aggregation, but are not in any way interdependent, so that the wholes are scarcely more than the sum of their parts.

This is, of course, a matter of how we define a multicellular organism, but I think any definition that excludes, for example, Eudorina, is not a very useful one.

This time, I’ll look at the rest of what Huxley had to say about the volvocine algae, most of which is about Volvox:

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J. S. Huxley part 1: Gonium

Julian Huxley was one of the biologists responsible for the merging of Mendelian genetics and Darwinian evolution in the early 20th century, the modern synthesis. His most influential work was Evolution: The Modern Synthesis, published in 1942. Thirty years earlier, though, he published a book on biological individuality, The Individual in the Animal Kingdom. Thankfully, the copyright on this book has expired, so it is now part of the public domain, and a scanned version is available for free in pdf and epub versions from Google.

Huxley Cover

Any book with Volvox on the cover can’t be all bad!

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The evolution of undifferentiated multicellularity: the Gonium genome

Blogging took a backseat to the wedding of two dear friends two weekends ago and to morel hunting last weekend, so I’m only now getting around to a post that should have been written weeks ago (I promised on April 22 that it would be out the following week). Last month, Erik Hanschen and colleagues published the Gonium pectorale genome, filling in some crucial bits of the transition to multicellular life in the volvocine algae. This was a big project, taking several years and involving over 20 authors from over a dozen institutions. The final paper is open access in Nature Communications.

I did post an effort to explain some aspects of the paper to the cdesign proponentsists at Evolution News and Views, who, by their own admission, failed to understand it (“After reading this paper, we’re none the wiser.”). I also complained of the science media’s tendency to refer to all algae as ‘pond scum.’ The lead author of the genome paper kindly followed up with a guest post addressing some of ENV‘s other misunderstandings, such as the purpose of model organisms in biology and the difference between ‘assertion’ and ‘evidence’. But now it’s time to dig into what the genome paper actually says.

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Please stop calling them pond scum

Gonium pectorale. Credit: Kansas State University.

Gonium pectorale. Credit: Kansas State University.

Yes, they live in ponds; no they don’t form any kind of scum. The press release from Kansas State on the Gonium genome paper, which is reprinted here, here, and here, is titled “Pond scum and the gene pool: One critical gene in green algae responsible for multicellular evolution, understanding of cancer origin.” Gonium forms planktonic colonies of (usually) 8, 16, or 32 cells that swim under their own power and exhibit phototaxis (they’ll swim toward a light source). They are not pond scum. ‘Algae’ and ‘pond scum’ are not synonyms, dig? Leaving aside the distinction between algae and cyanobacteria, calling Gonium pond scum is like saying pineapples are lemons (because both are fruits).

Also…cancer origin, really? You went there? The word ‘cancer’ does not appear in the paper except in the funding acknowledgements (Bradley Olson is partly funded by the KSU Johnson Cancer Center).

Heads I win; tails you lose: Evolution News & Views on Gonium, part 2: Model systems and gene duplication

Figure 2 from Hanschen et al. 2016. (a) Predicted number of genes in each phylostratum (PS1–PS9) for Chlamydomonas, Gonium and Volvox. (b) Heatmap of transcription factor abundance for all green algae. Significant over- (+) and under-representation (−) in colonial/multicellular lineages (Gonium and Volvox) is denoted (G test of independence, α=0.05). Rows are clustered (left), an accepted phylogeny is depicted (top). (c) Phylogenetic analysis of gene family evolution. Bars to the left and right of the vertical axis denote the lost and gained gene families respectively, relative to its parental node. (d) Venn diagram of the species distribution of Pfam A domains unique to the volvocine algae.

Figure 2 from Hanschen et al. 2016. (a) Predicted number of genes in each phylostratum (PS1–PS9) for Chlamydomonas, Gonium and Volvox. (b) Heatmap of transcription factor abundance for all green algae. Significant over- (+) and under-representation (−) in colonial/multicellular lineages (Gonium and Volvox) is denoted (G test of independence, α=0.05). Rows are clustered (left), an accepted phylogeny is depicted (top). (c) Phylogenetic analysis of gene family evolution. Bars to the left and right of the vertical axis denote the lost and gained gene families respectively, relative to its parental node. (d) Venn diagram of the species distribution of Pfam A domains unique to the volvocine algae.

Erik Hanschen, the lead author on the Gonium genome paper, is also an old friend of mine from when we were both in Michael Doebeli’s lab at the University of British Columbia. He kindly agreed to write a guest post responding to Evolution News and Views‘ misunderstandings of his paper. Everything below the fold was written by Erik:

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Heads I win; tails you lose: Evolution News & Views on Gonium, part 1

Figure 6 from Hanschen et al. 2016. Multicellularity hinges on the evolution of cell cycle regulation in a multicellular context with subsequent evolution of cellular differentiation (here, cell size-based) and increased body size.

Figure 6 from Hanschen et al. 2016. Multicellularity hinges on the evolution of cell cycle regulation in a multicellular context with subsequent evolution of cellular differentiation (here, cell size-based) and increased body size.

Remember how I said they’re prolific? Before I’ve even had a chance to write up my thoughts on the Gonium genome paperEvolution News & Views has already published theirs. The story has also been picked up by the Washington PostNew HistorianGenNews, and ScienceDaily (that last one looks like just a reprint of the press release from University of the Witwatersrand). By the way, the genome paper is open access, so you don’t need a subscription to see it for yourself.

We already know that cdesign proponentsists are not fans of research into the evolution of multicellularity, and that they have trouble understanding it. In an unsigned article on the Gonium genome at ENV, they admit that

After reading this paper, we’re none the wiser.

That’s too bad. I’m here to help.

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Gonium genome published

Figure 1 from Hanschen et al. 2016. (a) Evolution of cell cycle control (C), expanded ECM (E) and somatic cells (S) are denoted. (b) Micrographs of Chlamydomonas (green; scale bar, 10 μm), Gonium (blue; scale bar, 10 μm) and Volvox (black; scale bar, 25 μm) show morphological differences.

Figure 1 from Hanschen et al. 2016. (a) Evolution of cell cycle control (C), expanded ECM (E) and somatic cells (S) are denoted. (b) Micrographs of Chlamydomonas (green; scale bar, 10 μm), Gonium (blue; scale bar, 10 μm) and Volvox (black; scale bar, 25 μm) show morphological differences.

I haven’t read it yet and won’t have time today, but the Gonium pectorale genome paper just came out in Nature Communications! Erik Hanschen is the lead author, and the article is open access. I previously reported on Erik’s talk at Volvox 2015:

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