The sensory dorsal organs of crustaceans

The cuticle of crustaceans bears numerous organs, of which the functions of many are unknown. One of these, the sensory dorsal organ (SDO), is present in a wide diversity of taxa. Here we critically review the variability, ultrastructure, distribution, and possible function of this enigmatic cuticular organ. Previous data are complemented by new observations on larvae and adults of various malacostracans. The SDO is composed of four sensors arranged as the corners of a square, the centre of which is occupied by a gland. Pores or pegs surrounding this central complex may also form part of the organ. The arrangement and the external aspect of the five main elements varies greatly, but this apparently has little impact on their ultrastructural organisation. The sensors and the gland are associated with a particularly thin cuticle. Each sensor contains four outer dendritic segments and the central gland is made of a single large cell. It is not yet known what this large cell secretes. The SDO is innervated from the tritocerebrum and therefore belongs to the third cephalic segment. A similar organ, here called the posterior SDO, has been repeatedly observed more posteriorly on the carapace. It resembles the SDO but has a greater number of sensors (usually six, but up to ten) apparently associated with only two outer dendritic segments. The SDO and the posterior SDO are known in the Eumalacostraca, the Hoplocarida, and the Phyllocarida. Some branchiopods also possess a dorsal organ' resembling both the SDO and the ion-transporting organ more typical of this group. This may indicate a common origin for these two functionally distinct groups of organs. New observations on the posterior SDO support the hypothesis that the SDO and the posterior SDO are homologous to the lattice organ complexes of thecostracans. However, the relationship between the SDO and the dorsal cephalic hump of calanoid copepods remains unclear. No correlation can be demonstrated between the presence of a SDO and a particular ecological or biological trait. In fossils, the most convincing examples of SDO-like organs are found in some Late Cambrian arthropods from the Alum Shale of southern Sweden. They suggest that related organs might have been present in non-crustacean Cambrian arthropods. The distribution of the SDO and posterior SDO in extant and fossil crustaceans strongly suggests that these organs originated early in the history of the group, and are crucial to the functioning of these organisms. However, except for knowing that the sensors are chemoreceptors and that in a given organ a functional relationship probably exists between them and the gland, little is known about this function. The description of a SDO in freshwater carideans, which can be easily reared in a laboratory, opens the way for behavioural and physiological experiments to be undertaken that could prove crucial for the determination of this function.