Pathways to pluralism: Beckett Sterner on biological individuality, part 1

In grad school I wound up hanging around with John Pepper (yeah, Dr. Pepper) a good bit. I think I disagreed with him more than I agreed with him, sometimes to the point of exasperation, but conversations with him were never boring.


One of John’s most annoying refrains was “is it an organism?” I was studying (and still study) a group of algae for which this question can be genuinely confusing. Most people would say a Chlamydomonas cell is a single-celled organism, and most would agree that Volvox is a multicellular organism, but what about the four-celled species Tetrabaena? A four-celled organism or a collection of four single-celled organisms? What about an undifferentiated colony of 32 cells, such as Eudorina? Or Pleodorina, which is around the same size but with two cell types? Somewhere between a unicellular ancestor and Volvox, a new kind of individual emerged. Among the extant species*, where do we draw the line between organisms and groups of organisms, or can we (or should we) draw a line at all?

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

Van Leeuwenhoek 1700 Figure 5

Figure 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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Actin evolution in the Volvocales

Kato-Minoura Figure 1

Fig. 1 from Kato-Minoura et al. 2015: Genomic structure of volvocine actin and NAP genes. For comparison, previously identified sequences are also shown. Filled boxes, putative coding exons; open boxes, putative 5′ and 3′ untranslated regions. Intervening sequences are shown by solid lines. Intron positions are indicated by codon and phase numbers with reference to the three alpha-actins of vertebrates (377 amino acids) (Weber and Kabsch 1994). The conserved intron positions are linked with dotted lines. ATG, translation start codon; TAA or TGA, stop codon.

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African Volvox in Montana

Ninepipe Reservoir

Ninepipe Reservoir near Charlo, MT. Photo by Aeravi.

Last summer, I hosted Drs. Hisayoshi Nozaki, Noriko Ueki, Osami Misumi, and two graduate students from the University of Tokyo, Kayoko Yamamoto and Shota Yamashita, to collect volvocine algae from Montana lakes. To our surprise, we found a species of Volvox (V. capensis) that had previously only ever been found in South Africa! Dr. Nozaki’s team identified the algae collected in Ninepipe Reservoir based on their morphology and DNA sequencing. South Africa and Montana: this is about as disjunct as a distribution can be. Is Volvox capensis a master of long-distance dispersal? Is its distribution actually cosmopolitan, and if so, why hasn’t it ever been found anywhere else?

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Pierrick Bourrat responds

[I invited Pierrick Bourrat to respond to my two posts about his new paper and to comments to those posts. He kindly agreed, and he provided the following guest post, which I have edited only for formatting.]

First of all, I would like to thank Matthew Herron for his interest in my work and his invitation to respond to his posts. Also, I would like to thank Rick Michod and Deborah Shelton for their comments.

I will respond to several issues pointed out both in the posts and the comments.

About the usefulness of the export of fitness view of ETI: I agree that it is a useful way of thinking about it, as long as it is used as a heuristic. This means that I am not inclined to think that building models with the assumption that the fitness of a cell would have been 0 had it been in an environment with not social partners will be able to explain in some deep sense ETIs (and even more so the origin of fitness at some level). In his comment to Matthew’s first post, Rick Michod claims that I somehow confuse realized fitness from a more counterfactual notion of fitness.  Well, to be honest, I do not see how one could simulate (I do not mean ‘explain’) the evolution of a process if the variables in the model do not correspond to realized properties of the system. If I want to model a particular phenomenon, I ought to use variables and parameters that represent the target system and clearly, at least for me, this counterfactual notion of fitness does not represent any properties the cells have because they always have social partners. It is common to use expected rather than realized fitness in models, but this assumption is justified when we can assume that population are large and the environment is overall not fluctuating too much. With the counterfactual notion of fitness, aside from being useful for explaining the ETIs, I fail to see how it could be successfully integrated in models (by successfully, I mean how it could represent meaningfully the target system).

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Expression and form: Arash Kianianmomeni on gene regulation

Kianianmomeni Figure 1

Figure 1 from Kianianmomeni 2015. Gene regulatory mechanisms behind the evolution of multicellularity. Model illustrating the role of gene regulatory mechanisms in the evolution of multicellular Volvox from a Chlamydomonas-like ancestor.

Arash Kianianmomeni’s latest paper in Communicative & Integrative Biology addresses the possible roles of gene regulation and alternative splicing in the evolution of multicellularity and cellular differentiation (Kianianmomeni, A. 2015. Potential impact of gene regulatory mechanisms on the evolution of multicellularity in the volvocine algae. Commun. Integr. Biol., 37–41. doi 10.1080/19420889.2015.1017175). The article is an ‘Addendum’ to a 2014 study by Kianianmomeni and colleagues in BMC Genomics. Communicative & Integrative Biology often invites authors to write these addenda after they have published a (usually high impact) paper elsewhere, providing authors the opportunity to publish material that was not included in the original paper due to space limitations or because it was opinionated or speculative. I may address the BMC Genomics article in a future post, but right now there is more new volvocine research than I have time to write about (it should be an exciting Volvox meeting this summer!).

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Pierrick Bourrat on levels, time, and fitness, part 2: collective fitness

Last week, I posted some thoughts on Pierrick Bourrat’s new paper in Philosophy and Theory in Biology, focusing on his criticism of Rick Michod’s ‘export of fitness’ framework. This week, I’ll take a look at the second of Bourrat’s criticisms, regarding the transition from MLS1 to MLS2, as first defined by Damuth & Heisler, during a transition in individuality.
MLS1 and MLS2 refer to two different versions of MultiLevel Selection. As Bourrat describes it (and this is pretty much in line with other authors), fitness in MLS1 is defined in terms of the number of particles (or lower-level units, or cells) produced, while in MLS2 the fitnesses of the particles and collectives (or cells and multicellular organisms) are measured in different units. Cell-level fitness (for example) is defined in terms of the number of daughter cells, organism-level fitness is based on the number of daughter organisms. (As with last week’s post, I’ll generally stick to cells and organisms, though the principles apply equally to any two adjacent levels.

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Pleodorina study featured on NAI website

My new paper in Evolutionary Ecology Research is currently featured on the NASA Astrobiology Institute website (“Algae Fitness and Multicellular Life“). This was the final chapter of my Ph.D. dissertation, and it describes an artificial selection experiment using Pleodorina starrii. The paper is co-authored by my Ph.D. advisor, Rick Michod, and two (then) undergraduates, Susma Ghimire and Conner Vinikoor.
Pleodorina starrii

A 32-celled colony of Pleodorina starrii with 12 somatic cells.

Pleodorina is considered “partially differentiated,” meaning that some of its cells are of the ancestral, undifferentiated type (like those of Eudorina) and some are differentiated as somatic cells. These somatic cells never grow much, and they never divide to form daughter colonies.

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