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tunicates - apparent simplicity belies a complex past

Tunicates are more commonly known as 'sea squirts' - little blobby marine creatures that squirt water when you touch them (hence the name). We don't hear about them often, except perhaps when they make the news for all the wrong reasons. But from an evolutionary perspective they are fascinating little creatures - and it's largely due to their larvae.

As an aside: why do we call them tunicates? Because the body of the adult organism is enclosed in an outer sheath, aka a tunic. The majority of tunicate species belong to a group known as ascidians, which as adults live in shallow waters, attached to rocks or maritime structures (including boats). The remainder are planktonic & found out in the open ocean.

The larvae of many ascidians are free-swimming and, because of their body form, are often described as 'tadpole larvae'. (Some of the non-ascidian tunicates have adults with the same morphology.) These little animals have a number of features (shared with creatures such as the cephalochordate formerly known as Amphioxus) that link them with the chordates: a hollow dorsal nerve cord, a post-anal tail, a pharynx with slits in it (which feeds into the gut), and a living cartilaginous rod known as the notochord, against which the animal's muscles work. (The larvae, and the adults of some non-ascidian tunicates, are basically little swimming filtration units.)

In fact, because of their rather simple structure, tunicates have long been viewed as representing the likely common ancestor of both chordates (a group that includes us) and the slightly-more-complex cephalochordates like Amphioxus. However, a newly-published & fascinating article by Linda Holland (2016) looks at 

the highly derived body plans and life styles of the tunicate classes, their importance in the marine food web and their genomics [with an] emphasis ... on the impact of their especially rapid evolutionary rates on understanding how vertebrates evolved from their invertebrate ancestors.

It turns out that a genomic comparison, using nuclear genes from chordates, cephalochordates and tunicates, indicates that it's actually Amphioxus that sits at the base of this particular group. This in turn means that tunicates

have lost a lot of what the long extinct ancestral tunicate once possessed. 

This genomic work is fascinating on a number of levels. For example, the 'textbook wisdom' is only bacteria (ie Prokaryotes) have their genome organised into operons, where a single mRNA transcript contains several genes. But it turns out that tunicates, which have a rather small genome.

[have] a high percentage of genes in operons

something that Holland states they share with roundworms (nematodes) and some flatworms, which apparently also have "reduced genomes". In tunicates, it seems that among the genes that have been lost are some of the 'Hox' genes - genes that control the development and patterning of body form. 

I learned heaps of new things from this paper: tunicates are able to regenerate most of their bodies, for example (makes sense, I guess, as the sessile adult sea squirt can't exactly avoid being snacked on by predators). Apparently this is achieved by pluripotent stem cells in the animals' blood, though how it's done is still something of a mystery. And I had no idea at all that the animal's 'tunic'

contains cellulose, synthesized by a cellulose synthase that was evidently acquired in an ancestral tunicate by horizontal gene transfer from a bacterium. 

An animal that produces cellulose! Nature never ceases to surprise :)

L.Z.Holland (2016) Tunicates. Current Biology 26: 4 pR146–R152 DOI: http://dx.doi.org/10.1016/j.cub.2015.12.024

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