Vitellocytes and vitellogenesis in cestodes in relation to embryonic development, egg production and life cycle

Vitellocytes have two important functions in cestode embryogenesis: (1) formation of hard egg-shell (e.g. Pseudophyllidea) or a delicate capsule (e.g. Cyclophyllidea), and (2) supplying nutritive reserves for the developing embryos. During evolution any of these two functions can be reduced or intensified in different taxa depending on the type of their embryonic development, degree of ovoviviparity and life cycles. Within the Cestoda, there are three monozoic taxa with only one set of genital organs: Amphilinidea, Gyrocotylidea and Caryophyllidea. In these monozoic taxa and some polyzoic groups with well developed vitellaria (e.g. Pseudophyllidea, Trypanorhyncha) a single oocyte [=germocyte] and a large number of vitellocytes (up to 30) are enclosed within a thick, hardened egg-shell, forming a type of eggs typical for the basic pattern of Neodermata. Only one type of egg-shell enclosures, the so-called 'heterogeneous shell-globule vesicle' is common for the above mentioned cestode taxa. Each membrane-bounded vesicle of mature vitellocytes contains numerous electron-dense shell globules embedded in a translucent matrix. In free-living Neoophora and Monogenea there are two types of vesicles with dense granules; the second is considered to be proteinaceous reserve material. Within the Cestoda, the numbers of vitellocytes per germocyte are reduced in those taxa forming eggs of the 'Cyclophyllidean-type' (e.g. Cyclophyllidea, Tetraphyllidea, Pseudophyllidea). This is particularly evident in Cyclophyllidea; for example, in vitellocytes of Hymenolepis diminuta (Hymenolepididae) there are numerous vitelline granules of homogeneously electron-dense material; in Catenotaenia pusilla (Catenotaeniidae) there are three large, homogenous vitelline vesicles, while in Inermicapsifer madagascariensis (Anoplocephalidae) there is only one large vitelline vesicle, containing homogeneously electron-dense material, which occupies most of the vitelline cell volume. In this respect the Tetraphyllidea and Proteocephalidea, in forming eggs that lack a hard egg-shell, hold an intermediate position. A comparison of interrelationships which exist among types of vitellocytes, vitellogenesis, types of embryonic development, ovoviviparity and life cycles indicates parallelisms and analogies in adaptation to the parasitic way of life in different groups of cestodes. Knowledge on cestode vitellogenesis may also have an important applied aspect. Vitellocytes, due to their high metabolic rate, represent a very sensitive target for analysing effect of anthelminthic drugs upon the egg formation (ovicidal effects); rapid degeneration of vitellocytes is usually accompanied by a cessation of egg production.