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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Volvox inversion review

Alexey Desnitskiy from St. Petersburg State University has published a short review of the process of embryonic inversion in the genus Volvox. It is a translation, by the author, of his Russian-language paper in the journal Ontogenez (Desnitskiy, AG. 2018. Ontogenez 49:147-152). The article, in the Russian Journal of Developmental Biology, isn’t listed as open access, but it also doesn’t seem to be paywalled.

Inversion occurs during the development of all known species in the family Volvocaceae (Colemanosphaera, Eudorina, Pandorina, Platydorina, Pleodorina, Volvox, Volvulina, and Yamagishiella), where it serves to turn the embryo inside-out and get the flagella on the outer surface of the colony. The paper discusses the two distinct inversion processes found in different Volvox species:

…the inversion of “type A” and the inversion of “type B,” represented by the two species most thoroughly studied, respectively V. carteri f. nagariensis and V. globator (Hallmann, 2006; Höhn and Hallmann, 2011). The principal difference between these two types of inversion is that this process begins at the anterior pole of the embryo in the first case, while in its posterior hemisphere in the second case. Coordinated displacements of cells relative to the system of intercellular cytoplasmic bridges play, along with changes of the cell shape, an important role in the inversion process in embryos of both Volvox species. In V. globator, though, the spindle-shaped cells could be observed not in the entire embryo but only in the posterior hemisphere at the stage of its compression.

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A is for Algae

I got my copy of Jillian Freese’s A is for Algae earlier this week. Freese, a Ph.D. candidate at the University of Rhode Island, says the book is “Part birthday gift. Part #scicomm. Part stress relief.” It’s full of watercolor paintings of algae, mostly seaweeds but with some phytoplankton as well. Each species (one for each letter of the alphabet) is presented with its scientific name, usually a common name, habitat and biogeographic information, and some interesting factoids.A is for Algae

Warning: spoilers below the fold.

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Everything Flows

Everything Flows cover

Process philosophy has really just recently come on my radar, and I’m not sure what to make of it. I have written before that I don’t have a particularly strong background in philosophy, and so I’m hesitant to judge what I may not understand. At least some of the descriptions I’ve seen strike me as quasi-mystical word salads:

In short, a becoming actual entity prehends, or “feels,” not only other, past actual entities (which may be seen as the metaphysical basis for causality wherein one entity becomes part of another entity’s formation process), but also eternal objects (i.e., “pure possibilities”), which introduces novelty into the process. –Lukasz Lamza in Nature Alive – Essays on the Emergence and Evolution of Living Agents

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Waltzing Volvox

I can’t believe I haven’t already blogged about this, but if I have it isn’t turning up in my searches. Ravi Balasubramanian’s preprint about “flocking” behavior in Volvox barberi mentioned that

V[olvox] carteri is capable of using fluid forces created by flagellar beating to form waltzing pairs.

He’s referring to a 2009 paper by Knut Drescher and colleagues in Physical Review Letters. Drescher and colleagues analyzed the physics that cause Volvox colonies to enter a hydrodynamically bound state in which two or more spheroids orbit each other in close proximity:

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Alternative patterns of explanation for major transitions

The Major Transitions in Evolution Revisited

One reason to study green algae is because they can teach us something about the evolution of multicellularity. A number of related species in the Volvocalean family form a gradation of complexity between single-celled and simple multicellular organisms. The members of this family of algae differ in size, the number of cells they produce, and whether or not there is a split between germline and somatic cells. This split is thought to be central to understanding how a new level of individuality has evolved. — Calcott 2011, p. 39.

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Extreme variation in male Volvox carteri from Taiwan

Nozaki et al. 2018 Fig. 1 A-D

Figure 1 a-d from Nozaki et al. 2018. Light microscopy of asexual spheroids in Taiwanese strains of Volvox carteri f. nagariensis. a Surface view of a spheroid showing undivided gonidia (G). 2016‐tw‐nuk‐6‐1. b Optical section of a spheroid in (a) with gonidia (G). c Surface view of spheroid. Note no cytoplasmic bridges between somatic cells. 2016‐tw‐nuk‐6‐1. d Surface view of spheroid showing individual sheaths of the gelatinous matrix. Stained with methylene blue. 2016‐tw‐nuk‐8‐1. e Optical section of gonidium. 2016‐tw‐nuk‐6‐1. f, g Pre‐inversion plakea or embryo (E) showing gonidia (G) of the next generation outside. 2016‐tw‐nuk‐8‐1.

Most of what we know about the developmental genetics of Volvox comes from the Eve strain of Volvox carteri forma nagariensis, which was collected by Richard Starr from Kobe, Japan in 1967. Eve is the strain that David Kirk and colleagues used for most of their experiments and from which most of the important developmental mutants are derived.

It’s natural, then, to think that Eve is representative of V. carteri f. nagariensis and that what’s true for Eve is generally true for this forma. Recent work from Hisayoshi Nozaki and colleagues shows that, at least in one respect, this is a bad approximation.

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Volvox on Micropia

Volvox (Micropia)

Image from www.micropia.nl/en/discover/microbiology/volvox/

Micropia, the museum of microbes in Amsterdam, has a page devoted to Volvox:

Ponds and ditches are not only home to unicellular green algae, but also to multicellular forms.

Some ‘colonies’ are nothing more than a mass of single cells all doing exactly the same thing, but with the spherical volvox it’s a slightly different story. Here different cells have specialised and work together. All the cells are located on the outside of the sphere. There are cells with flagella (whip-like hairs) to help the colony move around and cells which are responsible for reproduction.

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