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Metazoa |--Porifera (paraphyletic?) `--+--Radiata `--Bilateria |--Deuterostomia `--Protostomia |--Ecdysozoa `--Spiralia
The Problem of Middle Earth
Trees in the Entwood
"Where Many Paths and Errands Meet"
This category includes all Metazoa with bilateral symmetry (having a definite front and rear, and left and right body surfaces), either in their adult stage or (in the case of types with secondary radial symmetry, such as echinoderms) in the larval stage. Many bilaterian animals exhibit cephalization, which is the evolutionary trend toward concentration of sensory structures, mouth, and nerve ganglia, at the anterior end of the body. This body type is related to the further elaboration of hox genes
All bilaterians are triploblastic, which means they develop three germ layers during embryonic development. The three germ layers are
Note that only Bilateria have a mesoderm.
Bilateria is defined by reference to a hypothetical organism, Urbilateria, who was the last common ancestor of Deuterostomia (echinoderms and the various chordate phyla) and Protostomia (all other "higher" animals). Classically, these two groups were said to differ in a number of respects. Some of these characteristics have been challenged recently, but they remain generally true:
Update (April 2007): A number of papers published in the last few years have made it clear that the whole traditional concept of Bilateria is probably mistaken. Bilateria supposedly have three unique features: (1) bilateral symmetry, (2) hox gene patterning of the anteroposterior axis, and (3) mesoderm. It turns out that developing cnidarians clearly have (1), probably have (2), and may even have (3). It seems even more likely now that the Cnidaria are paraphyletic. That is, all of us bilaterian animals are just jellyfish who happened to use a particular type bilateral symmetry and used the hom genes in a particular way.
This may have important phylogenetic implications. We cannot assume that any bilateral organism with a couple of the usual hom genes is inside the crown group of Bilateria. Consequently, the predictions we made last year (see end of this essay) about Metazoan phylogeny are looking good.
Most of this is covered in Brooke & Holland (2003), which we cited -- but obviously didn't pay enough attention to. To see how far things have progressed at this point, see Ryan et al. (2007). ATW070404
One important difference involves the cleavage patterns; the division of cells in the early embryo. Protostomes are characterized by a spiral (the planes of cell division are diagonal to the vertical axis of the embryo) and determinant (the goal of each embryonic cell is established very early) cleavage. Deuterostomes undergo radial (parallel or perpendicular to the vertical axis of the embryo) and indeterminate cleavage (each early embryonic cell retains the capacity to develop into a complete embryo if isolated from other cells). But see, Halanych (2004); Hejnol & Schnabel (2004).
There is a strange symmetry in the fate of the blastopore (the first opening of the archenteron which forms during gastrulation.). In Protostomes ("mouth first") the blastopore forms the mouth; in Deuterostomes ("mouth second") - blastopore forms the anus. (see following diagram) .
Another difference involves how the mesoderm surrounds the body cavity. It can either pouch off or split entirely. In Schizocoela or Protostomia, it splits. In the Enterocoela or Deuterostomia, the mesoderm (or archenteron, the original gut) pouches off
Bilateria Hatschek, 1888 : fish > jellyfish
Range: Fr Ediacaran
Phylogeny: Eumetazoa : Cnidara + * : Protostomia + Deuterostomia
Characters: Primary bilateral symmetry, occaisonally secondarily modified to pentameral or radial, organ-system grade of organization, most triploblastic with well-developed mesoderm of endodermal origin, most with body cavity other than the digestive cavity, anus typically present.
Comments: For simplicity's sake, minor and problematic phyla have been excluded from the Phylogeny line; these will be added at some later point. The Bilateria evolved from a paraphyletic Radiata; early bilateral phylogeny remains obscure, MAK120423
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this page MAK120423 (from bilateria.htm)