Origins of the sexes: isogamy and anisogamy

Sex didn’t always involve males and females. I know it still isn’t always between males and females, but that’s not what I mean. I mean that there was a time when sex was happening, but there were no males and females. Sex existed before males and females, and many species are still doing it without them.

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Initiation of cell division in Chlamydomonas

Deborah Shelton and colleagues have published a new article arguing that the reigning model of cell division initiation in Chlamydomonas reinhardtii needs to be revised [full disclosure: Dr. Shelton and I were labmates in Rick Michod’s lab at the University of Arizona]. The evolution of multicellularity almost certainly involved changes in cell cycle regulation; for example, there is good evidence that changes to the cell cycle regulator retinoblastoma were involved in the initial transition to multicellular life in the volvocine algae. So understanding cell cycle regulation is vital for understanding the evolution of multicellularity.

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Pleodorina inversion

Stephanie Höhn and Armin Hallmann have published a detailed study of the developmental process of inversion in Pleodorina californicaPleodorina is one of the two genera we usually refer to as ‘partially differentiated’ (the other is Astrephomene), meaning that some of their cells are specialized for motility and never reproduce (soma) and some perform both motility and reproductive functions. P. californica is pretty big, up to about 1/3 of a millimeter, easily visible to the naked eye (though you’d need better vision than mine to make out any details).

Stephanie Höhn sampling a pond near Cambridge University during the Volvox 2015 meeting.

Stephanie Höhn sampling a pond near Cambridge University during the Volvox 2015 meeting.

Like all members of the family Volvocaceae, P. californica undergoes complete inversion during development:

After the completion of the cell division phase and before inversion, the embryos of Gonium, Pandorina, Eudorina and Pleodorina consist of a bowl-shaped cell sheet, whereas the embryonic cells of Volvox form a spherical cell sheet. With exception of the genus Astrephomene, all multicellular volvocine embryos face the same “problem”: the flagellar ends of all the cells point toward the interior of the bowl-shaped or spherical cell sheet rather than to the exterior, where they need to be later to function during locomotion. [References removed]

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Multicellularity rundown

Too many papers, not enough time: each of these deserves a deep dive, but my list just keeps getting longer, so I’m going to have to settle for a quick survey instead. To give you an idea of what I’m up against, these papers were all published (or posted to bioRxiv) in July and August, 2016. By the time I could possibly write full-length posts about them all, there would probably be ten more!

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Retrogenes in Volvox and Chlamy

The evolution of multicellularity in the volvocine algae appears to have happened primarily through co-option of existing genes for new functions. For example, the initial transition from a unicellular life cycle to a simple multicellular one involved the retinoblastoma gene, as Hanschen and colleagues elegantly demonstrated (see “The evolution of undifferentiated multicellularity: the Gonium genome“). A Volvox gene involved in cellular differentiation, regA, was likely co-opted from an ancestral role in environmental sensing, and a similar origin appears to explain the use of cyclic AMP for the signaling that causes multicellular aggregation in cellular slime molds (see “Volvox 2015: evolution“). 

Some of the changes leading to complex multicellularity, though, clearly did involve new genes. Two gene families involved in building the extracellular matrix that makes up most of a Volvox colony, the pherophorins and metalloproteinases, have undergone multiple duplication events leading to greatly expanded gene families (see “Heads I win; tails you lose: Evolution News & Views on Gonium, part 2“). One mechanism by which genes are duplicated is retroposition, in which a messenger RNA is reverse transcribed into DNA and inserted into the genome:

Fig S1A from Jakalski et al. 2016. Basic mechanism of retroposition. DNA is transcribed into a pre-mRNA by RNA polymerase, introns are spliced out, and a poly(A) tail is added to the 3′ end, resulting in a mature messenger RNA. The mRNA is then reverse-transcribed to DNA and inserted into a new genomic location.

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Graduate student position in the Nedelcu lab

If you’re a fan of Volvox and the volvocine algae and have recently received an undergraduate degree in biology or a related field, now’s your chance to get serious about studying them. Aurora Nedelcu is looking for a graduate student to join her lab at the University of New Brunswick. Professor Nedelcu is a major player in the Volvox community, having published foundational papers on diverse aspects of volvocine biology and organized the first two international Volvox meetings. This is a great opportunity to join a vibrant and growing research community:

A graduate student position is available in the laboratory of Aurora Nedelcu, in the Department of Biology at the University of New Brunswick, Fredericton, CANADA. Research in our laboratory is directed towards understanding general, fundamental issues in evolution – such as the evolution of multicellularity, development, cell differentiation, sex, programmed cell death, altruism.  Our research is rooted in the framework of transitions in individuality and evolution of complexity (at a conceptual level), and of cellular responses to stress (at a more mechanistic level).  The experimental model-system we are currently using is the green algal group, Volvocales (see our Volvocales Information Project; Highly motivated students with interests in either theoretical/genomics or experimental/molecular approaches, and previous research experience are encouraged to apply. Interested applicants should e-mail a CV, summary of research experience and interests, unofficial transcripts, and contact information for three referees to

Applicants should meet the minimum requirements for acceptance in the Biology Department Graduate Program (see

I’m going to Cyprus!


I’m heading to Limassol, Cyprus at the end of September to talk about Volvox morphology and evolution. Phycomorph is a European group studying seaweed development and reproduction, with a large focus on cultivation. I have an extra day after the meeting, so hopefully I’ll get to explore a bit.

The organizers were initially worried about the high cost of transportation from Missoula, but I had good news for them: I won’t be flying from Missoula but from Atlanta, which is (seriously) half as expensive. The timing of the flights is a bit unfortunate, though, so I’ll have a couple of very long layovers in Heathrow (17 and 20 hours).


The preliminary list of confirmed speakers is:

Phycomorph speakers


Problems with major transitions: Maureen O’Malley & Russell Powell respond

The Great Oxidation Event by Adelle Schemm.

The Great Oxidation Event by Adelle Schemm.

In a recent series of posts, I reviewed Maureen O’Malley and Russell Powell’s paper in Biology and Philosophy, “Major Problems in Evolutionary Transitions: How a Metabolic Perspective Can Enrich our Understanding of Macroevolution.” Although they made several good points, I thought that some of their criticisms were off the mark and that their proposed solution to the real and perceived problems with the major transitions framework was unsatisfying.

Drs. O’Malley and Powell are both heavy hitters in the philosophy of biology, and as I usually do when I dig deeply into someone else’s paper, I invited them to respond to my criticisms. They kindly provided a thoughtful rebuttal and permitted me to post it here. I’ll have more to say later, but for now I’ll just say that they make some good points and (most importantly) fairly represent my arguments. As usual for guest posts, I have made no edits to the content of their response, only formatted and added links:

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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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