Image from www.twoheadedquarter.net.
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.
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; http://www.unbf.ca/vip). 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 anedelcu@unb.ca.
Applicants should meet the minimum requirements for acceptance in the Biology Department Graduate Program (see http://www2.unb.ca/biology/Degree_Info/Graduate.html).
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:
Maureen O’Malley and Russell Powell say that the major transitions framework is in need of repair. They have a point, or rather several good points. I have looked at their criticisms of three different versions (the original framework as laid out in the book by John Maynard Smith and Eörs Szathmáry, Rick Michod’s ‘evolutionary transitions in individuality‘ framework, and Szathmáry’s revised ‘Major Transitions 2.0‘). But what is their proposed fix, and will it have the intended effect?
Figure 4 from O’Malley and Powell 2016. Two major aeons of evolution (modified from Falkowski 2006). ‘Gya’ stands for ‘billion years ago’; the date for the origin of photosynthesis may need to be pushed back (see Crowe et al. 2013).
One of the cool things about studying the so-called major transitions is that they are as interesting to philosophers of science as to biologists. So you really can’t help being exposed to the philosophy of science literature, and many (maybe most) biologists in the field cross the lines at least occasionally. I’ve been to both, and I’m here to tell you that philosophy conferences are more fun than biology conferences.
Last time, I briefly summarized the various forms of the major transitions framework and addressed one of O’Malley and Powell‘s criticisms, that the framework is progressivist. Now I’d like to look at their other two problems: lack of unity and missing events. By and large, I agree with these points, although there are some caveats I’d like to point out. Next time, I’ll consider their proposed solution, which I’m afraid I don’t find helpful.
Disunity is actually O’Malley and Powell’s first criticism, but it will be a bit more complicated than progressivism to address, and I was short on time on part 1. Essentially, they are arguing that the major transitions are not a natural kind, philosophese for groupings that belong together because of some fundamental commonality, as opposed to more arbitrary groupings whose members are only superficially similar. So what are the transitions? Here’s the list from the book:
Table 1.2 from Maynard Smith J, Szathmáry E (1995) The Major Transitions in Evolution. Oxford University Press, Oxford.