Choanoflagellates with inversion

Salpingoeca rosetta

Figure 1A from Dayel et al. 2011. Spherical colony of Salpingoeca rosetta. Scale bar = 5 μm.

The closest (known) living relatives of animals are a group of unicellular or colonial filter-feeders known as choanoflagellates. Much of what we know about the evolution of multicellularity in animals comes from comparisons with choanoflagellates. For example, many of the gene families involved in multicellular development in animals, and previously thought to be unique to animals, have turned out to be present in choanoflagellates as well, suggesting that these gene families were present in animal ancestors before they evolved multicellularity. Some multicellular choanoflagellates have even been shown to have differentiated cell types (Laundon et al. 2019):

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Upcoming talks, and some system maintenance

Chlamydomonas colonies from the predation experiment.

Chlamydomonas colonies from the predation experiment.

I’ll be giving a couple of talks on experimental evolution of multicellularity in the next couple of weeks:

  1. University of Georgia Department of Cellular Biology, Tuesday, September 11, 11:00 a.m. in Biological Sciences 404A
  2. Donald Danforth Plant Science Center, Friday, September 20, time and place TBD

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Fungi are weird

I think about the evolution of multicellularity a lot, and I talk about it with colleagues. One of the things we talk about is what general principles we can infer from the many independent origins of multicellularity, for example in land plants, animals, red algae, brown algae, green algae, and fungi. Those are the groups that have evolved what we might call complex multicellularity, and one of the things we notice is that they all develop clonally; that is, they start out as a single cell, and when that cell divides, the daughter cells stick together. We notice that complex multicellularity has never evolved in species with aggregative development, when free-living cells come together to form a multicellular body, as they do in cellular slime molds and myxobacteria. Some aggregative developers have evolved a couple of different cell types, but all of the groups that have reached higher degrees of complexity develop by cell division and the products of cell division staying together. All, that is, except for fungi. Fungi are weird.

Fungi don’t really develop clonally in the way I’ve described, but they don’t really not develop clonally either. That’s because their cells don’t divide in the way we’re used to thinking about, through repeated rounds of mitosis. In mitosis, duplication of the genome is coupled to cell division: the chromosomes duplicate, they move to either end of the cell, then the cell divides. The chromosomes double, then they halve, so the daughter cells end up with the same number as the mother cell. That’s not how it works in fungi. Instead, they form filaments called hyphae (singular hypha) that grow at the tip. In some cases, partitions called septa (singular septum) form behind the growing tip, dividing the hyphae into individual cells. In some cases, no septa form, and each hypha is effectively one long, skinny cell with lots of nuclei (this is called a coenocyte).

So fungi don’t really develop by repeated rounds of cell division in the same sense that animals, plants, etc. do. Hyphae just grow, and they are divided into cells as sort of an afterthought, if they are divided into cells at all. Fungi with coenocytic (or aseptate) hyphae aren’t really even multicellular in the same sense as plants and animals are. Different people have different qualifications for what counts as multicellular, but it’s a stretch to call something multicellular that doesn’t have multiple cells. Fungi are weird.

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Debunked by the Institute for Creation Research

Folks, it’s been fun. I feel like I had a pretty good run as a scientist. I met some amazing people, went to beautiful places, and learned things I never would have imagined (Hodgkinia, WTF?!). With all my frustrations and failures, I’ve never once regretted going back to school and becoming a biologist. But now I need to close the door on all of that and find a new way to make a living.

See, the main project I’ve been working on for the last six years, the one that was supported by a NASA postdoctoral fellowship, and that just came out in Scientific Reports, has been debunked:

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Postdoctoral fellowship in the evolution of multicellularity

Thompson lab

Image from https://thethompsonlab.wordpress.com/.

The Thompson lab at University College London is looking for a postdoctoral researcher to study the evolution of multicellularity:

The Thompson lab, based at University College London, is seeking a Research Fellow to work on understanding how gene network heterogeneity affects the evolution of multicellular development.

Recently, we found that cell-cell variation in cell cycle position facilitates symmetry breaking during development, as it primes cells to respond to different differentiation cues (Gruenheit et al, Developmental Cell, 2018).

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Are the multicellular volvocine algae monophyletic?

One of the strengths of the volvocine algae as a model system is that they span a range of sizes and degrees of complexity. Sizes range from tens of microns to a couple of millimeters, cell numbers range from one to 50,000 or so, some species do and some don’t have cellular differentiation, and some do and some don’t undergo inversion during development. This variation makes the volvocine algae ripe for comparative analyses, which I and many others have done. It also allows many of the intermediate steps between unicellular and complex multicellular life to be identified, as David Kirk did in his “twelve-step” paper.

The volvocine algae have clearly taken some of those steps more than once. Cellular differentiation, for example, has evolved at least three times, in the genus Astrephomene, in the so-called Volvox section Volvox (a.k.a. Euvolvox), and in the lineage that includes Pleodorina and the other Volvox species. One thing they seem to have only done once, though, is to evolve multicellularity itself.

There have been dozens of studies addressing the evolutionary relationships among various species of volvocine algae. Most have been from Hisayoshi Nozaki’s lab, though I and many others have weighed in as well. Nearly all of them, at least those that address the topic, agree that the three families that make up the multicellular volvocine algae–the Tetrabaenaceae, Goniaceae, and Volvocaceae–uniquely descend from a common ancestor. In other words, the multicellular volvocine algae are monophyletic.

Three important cladistic terms are used to summarize the evolutionary relationships among a group of species. If all of the members of the group descend from a common ancestor, and nothing else descends from that ancestor, the group is called monophyletic. Mammals, for example, are monophyletic. A monophyletic group is also called a clade. If all group members are descended from a common ancestor, but so are some non-group members, the group is called paraphyletic. Reptiles, for example, are paraphyletic, because there is no clade that includes all reptiles that doesn’t also include birds. The word ‘paraphyletic’ should nearly always be followed by ‘with respect to’: reptiles are paraphyletic with respect to birds.

The bottom of the barrel, in terms of evolutionary relationships, is polyphyly. A group is considered polyphyletic if its members don’t share a recent common ancestor at all, in other words, if they have multiple evolutionary origins. Flying animals are polyphyletic. Algae are polyphyletic. The genus Volvox is polyphyletic. Polyphyletic taxa are the scum of the phylogenetic Earth. Telling a taxonomist that a group she has named is polyphyletic is a deadly insult.

The prevailing view of volvocine evolutionary relationships is that the family Volvocaceae is sister to the Goniaceae (that is, each is the other’s closest relative), and the Tetrabaenaceae are sister to the Volvocaceae + Goniaceae. Two new papers infer relationships among volvocine algae and their unicellular relatives, and one of them challenges the view of multicellular monophyly.

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Ulvophyte multicellularity: the sea lettuce genome

Ulva

Sea lettuce (Ulva sp.), Jericho Beach, Vancouver, BC, February 28, 2011.

David Kirk called the Chlorophyte green algae “master colony-formers” because multicellularity has evolved so many times within this class:

Although members of most chlorophycean genera and species are unicellular flagellates, multicellular forms are present in 9 of the 11 chlorophycean orders (Melkonian 1990). Multicellularity is believed to have arisen independently in each of these orders, and in some orders more than once.

In contrast, multicellularity has probably only evolved once or twice in the probable sister group of the Chlorophyceae, the Ulvophyceae. So when numbers like 25 get thrown around for the number of times multicellularity has evolved, something like half of those times were in the green algae.

We know a lot less about how multicellularity evolved in the Ulvophyceae than we do in the volvocine algae within the Chlorophyceae. A big step forward in understanding ulvophyte multicellularity happened last week, though, with the publication of the Ulva mutabilis genome.

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Uncommon Descent on Elizabeth Pennisi’s Science article

Two-headed quarter

Image from www.twoheadedquarter.net.

Yesterday, I ran a bit long about Elizabeth Pennisi’s new article in Science, “The momentous transition to multicellular life may not have been so hard after all.” I’m not the only one who noticed it, though; Uncommon Descent also commented (“At Science: Maybe the transition from single cells to multicellular life wasn’t that hard?“). There’s not much to it, just a longish quote from the article followed by this:

So at the basic level, there is a program that adapts single cells to multicellularity? Yes, that certainly makes multicellularity easier and even swifter but it also make traditional Darwinian explanations sound ever more stretched.

So if the evolution of multicellularity is easy, that’s evidence against “traditional Darwinian explanations.” Remember “Heads I win, tails you lose“?

…if multicellularity is really complicated, that’s evidence for intelligent design. But if multicellularity is really simple, that’s evidence for intelligent design.

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