Segmentation (biology)

Segmentation is a difficult process to satisfactorily define. Many taxa (for example the molluscs) have some form of serial repetition in their units but are not conventionally thought of as segmented. Segmented animals are those considered to have organs that were repeated, or to have a body composed of self-similar units, but usually it is the parts of an organism that are referred to as being segmented. ==Embryology==

'', a millipede with 170 segments and 662 legs Segmentation in animals typically falls into three types, characteristic of different arthropods, vertebrates, and annelids. Arthropods such as the fruit fly form segments from a field of equivalent cells based on transcription factor gradients. Vertebrates like the zebrafish use oscillating gene expression to define segments known as somites. Annelids such as the leech use smaller blast cells budded off from large teloblast cells to define segments. Arthropods s in the body segments of different groups of arthropod, as traced by evolutionary developmental biology. The Hox genes 7, 8, and 9 correspond in these groups but are shifted (by heterochrony) by up to three segments. Segments with maxillipeds have Hox gene 7. Fossil trilobites probably had three body regions, each with a unique combination of Hox genes. Although Drosophila segmentation is not representative of the arthropod phylum in general, it is the most highly studied. Early screens to identify genes involved in cuticle development led to the discovery of a class of genes that was necessary for proper segmentation of the Drosophila embryo. To properly segment the Drosophila embryo, the anterior-posterior axis is defined by maternally supplied transcripts giving rise to gradients of these proteins. This gradient then defines the expression pattern for gap genes, which set up the boundaries between the different segments. The gradients produced from gap gene expression then define the expression pattern for the pair-rule genes. Finally, the number of segments within the embryo is defined by the number of divisions and blast cells. Within the annelids, as with the arthropods, the body wall, nervous system, kidneys, muscles and body cavity are generally segmented. However, this is not true for all of the traits all of the time: many lack segmentation in the body wall, coelom and musculature. The interaction of other signaling molecules, such as myogenic regulatory factors, with this gradient promotes the development of other structures, such as muscles, across the basic segments. Lower vertebrates such as zebrafish do not require retinoic acid repression of caudal Fgf8 for somitogenesis due to differences in gastrulation and neuromesodermal progenitor function compared to higher vertebrates. Other taxa In other taxa, there is some evidence of segmentation in some organs, but this segmentation is not pervasive to the full list of organs mentioned above for arthropods and annelids. One might think of the serially repeated units in many Cycloneuralia, or the segmented body armature of the chitons (which is not accompanied by a segmented coelom). ==Origin==

Segmentation can be seen as originating in two ways. To caricature, the 'amplification' pathway would involve a single-segment ancestral organism becoming segmented by repeating itself. This seems implausible, and the 'parcellization' framework is generally preferred – where existing organization of organ systems is 'formalized' from loosely defined packets into more rigid segments. As such, organisms with a loosely defined metamerism, whether internal (as some molluscs) or external (as onychophora), can be seen as 'precursors' to eusegmented organisms such as annelids or arthropods. ==See also==