Tag Archives: Trichoplax adhaerens

Who came first? The comb or the sponge?

by Piter Kehoma Boll

The endless question is here again, but this time it appears to be settled. What animal group is the earliest of all? Who came first?

It is clear that there are five animal lineages that are usually regarded as monophyletic: sponges, placozoans, comb jellies, cnidarians and bilaterians. Let’s take a brief look at each of them:

Sponges (phylum Porifera) are always sessile, i.e., they do not move and are fixed to the substrate. They have a very simple anatomical structure. Their body is consisted of a kind of tube, having a large internal cavity and two layers of cells, an outer one and an inner one around the cavity. There are several small openings connecting the cavity to the outside, called pores, and one or more large cavities, called oscula (singular: osculum). Between the two cell layers there is a jelly-like mesohyl containing unspecialized cells, as well as the skeleton structures, including fibers of spongine and spicules of calcium carbonate or silica. Some species also secrete an outer calcium carbonate skeleton over which the organic part grows. Sponges lack muscles, nervous system, excretory system or any other kind of system. They simply live by beating the flagella of the choanocytes (the cells of the inner layer), creating a water flow entering through the pores and exiting through the osculum. The choanocytes capture organic particles in the water and ingest them by phagocytosis. All sponge cells can change from one type to another and migrate from one layer to another, so there are no true tissues.

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Body structures found in sponges. Picture by Philip Chalmers.*

Placozoans (phylum Placozoa) are even simpler than sponges, but they actually have true tissues. They are flat amoeboid organisms with two layers of epithelium, one dorsal and one ventral, and a thin layer of stellate cells. The ventral cell layer is slightly concave and appears to be homologous to the endoderm (the “gut” layer) of other animals, while the upper layer is homologous to the ectoderm (the “skin” layer).

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Trichoplax adhaerens, the only species currently in the phylum Placozoa. Photo by Bernd Schierwater.**

Comb jellies (phylum Ctenophora) resemble jellyfishes, but a closer look reveals many differences. Externally they have an epidermis composed by two layers, an outer one that contains sensory cells, mucus-secreting cells and some specialized cells, like colloblasts that help capturing prey and cells containing multiple cilia used in locomotion, and an inner layer with a nerve net and muscle-like cells. They have a true mouth that leads to a pharynx and a stomach. From the stomach, a system os channels distribute the nutrients along the body. Opposite to the mouth there is a small anal pore that may excrete small unwanted particles, although most of the rejected material is expelled through the mouth. There is a layer of jelly-like material (mesoglea) between the gut and the epidermis.

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The comb jelly Bathocyroe fosteri.

Cnidarians (phylum Cnidaria) have a structure similar to comb jellies, but not as complex. They also have an outer epidermis, but this is composed by a single layer of cells, and a sac-like gut surrounded by epthelial cells (gastrodermis), as well as a mesoglea between the two. Around the mouth there is one or two sets of tentacles. The most distinguishing feature of cnidarians is the presence of harpoon-like nettle cells, the cnidocytes, which are used as a defense mechanism and to help subdue prey.

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Body structure of a cnidarian (jellyfish). Picture by Mariana Ruiz Villarreal.

Bilaterians (clade Bilateria) includes all other animals. They are far more complex and are characterized by a bilateral body, cephalization (they have heads) and three main cell layers, the ectoderm, which originates the epidermis and the nervous system, the mesoderm, which give rise to muscles and blood cells, and the endoderm, which develops into the digestive and endocrine systems.

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Basic bilaterian structure.

Traditionally, sponges were always seen as the most primitive animals due to their lack of true tissues, muscular cells, nervous cells and all that stuff. However, some recent molecular studies have put the comb jellies as the most primitive animals. This was highly unexpected, as comb jellies are far more complex than sponges and placozoans, which would suggest that muscles and a nervous system evolved twice in the animal kingdom, or that sponges are some weird simplification of a more complex ancestor, which would be very hard to explain. The nervous system of comb jellies is indeed quite unusual, but not so much that it needs an independent origin.

However, now things appear to be settled. A study published this month on Current Biology by Simion et al. reconstructed a phylogenetic tree using 1719 genes of 97 animal species, and applying new and more congruent methods. With this more refined dataset, they recovered the classical reconstruction that puts sponges at the base of the animal tree, a more plausible scenario after all.

But why other studies have found comb jellies as the most basal group? Well, it seeems that comb jellies have unusually high substitution rates, meaning that their genes evolve faster. This leads to a problem called “long branch attraction” in phylogenetic reconstructions. As DNA has only four different nucleobases, namely adenine, guanine, cytosine and thymine, each one can only mutate into one of the other three. When mutations occur very often, they may go back to what they were in long lost ancestor, leading to misinterpretations in the evolutionary relationships. That seems to be what happens with comb jellies.

So, it seems that after all the sponge indeed came first.

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References and further reading:

Borowiec ML, Lee EK, Chiu JC, & Plachetzki DC 2015. Extracting phylogenetic signal and accounting for bias in whole-genome data sets supports the Ctenophora as sister to remaining Metazoa. BMC Genomics 16: 987. DOI: 10.1186/s12864-015-2146-4

Littlewood DTJ 2017. Animal Evolution: Last Word on Sponges-First? Current Biology 27: R259–R261. DOI: 10.1016/j.cub.2017.02.042

Simion P, Philippe H, Baurain D, Jager M, Richter DJ, Di Franco A, Roure B, Satoh N, Quéinnec É, Ereskovsky A, Lapébie P, Corre E, Delsuc F, King N, Wörheide G, & Manuel M 2017. A Large and Consistent Phylogenomic Dataset Supports Sponges as the Sister Group to All Other Animals. Current Biology 27: 958–967. DOI: 10.1016/j.cub.2017.02.031

Wallberg A, Thollesson M, Farris JS, & Jondelius U 2004. The phylogenetic position of the comb jellies (Ctenophora) and the importance of taxonomic sampling. Cladistics 20: 558–578. DOI: 10.1111/j.1096-0031.2004.00041.x
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Filed under cnidarians, Evolution, sponges, Zoology