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TitleSpermiogenesis, sperm ultrastructure and reproductive tract morphology in cicadas : implications for systematic relationships
AuthorChawanji, A.S.
SubjectSubjects to be assigned
AbstractSperm structure in five species of cicadine cicadas (Albanycada albigera, Azanicada zuluensis, Platypleura capensis, P. hirtipennis and Pycna semiclara) and five species of cicadettine cicadas (Melampsalta leucoptera, Quintilia walkeri, Stagira simplex, Xosopsaltria thunbergi and Monomatapa matoposa) was investigated by light and electron microscopy. In addition, spermiogenesis in cicadas was described; the information was derived from two cicadettines (Diceroprocta biconica and M. matoposa) and three cicadines (Kongota punctigera, P. capensis and P. semiclara). Mature spermatozoa of all species investigated are elongate and filiform, consisting of three distinct regions: the head (acrosome and nucleus), mid-piece and tail. All species produce more than one discrete length of nucleated, motile sperm, a form of sperm polymorphism termed polymegaly. Polymegaly is expressed in three ways; sperm have uni-, bi- or trimodal nucleus and tail lengths. Besides the differences in length, there are also notable differences in the size of nuclei. The anterior parts of sperm heads are embedded in an elongate homogenous matrix forming spermatodesmata. The conical acrosome is deeply invaginated posteriorly, and sits on top of the nucleus. The acrosomal contents are differentiated internally with a tubular substructure and a subacrosomal space. The anterior of the nucleus intrudes into the posterior section of the subacrosomal space. Anteriorly the acrosome is laterally flattened; posteriorly it extends as two tubular processes on either side of the nucleus that gradually decrease in diameter. The homogenously electron-dense nucleus is pointed anteriorly and is generally cylindrical, although posteriorly there is a lateral invagination that extends part-way along the nucleus. This invagination houses fine granular material of the putative centriolar adjunct which does not form in close proximity to the centriole and hence may not be a true centriolar adjunct. The lamellate disposition of the centriolar adjunct material within the sperm-midpiece of cicadettine cicadas is distinct, and separates these cicadas from their cicadine counterparts in which the centriolar adjunct material is non-lamellate. Vesicle-like elements that are associated with both the posterior nucleus and the centriolar adjunct are also found within the invagination. Immediately posterior of and adjoining the centriolar adjunct is a pair of mitochondrial derivatives that are elongated and extend for almost the entire length of the tail. Except for size the architecture of short and long spermatozoa is generally similar in all species. The absence of accessory bodies in cicada sperm suggests that within the Cicadomorpha, the families Cicadidae and Cercopidae are closely related. Only long nuclei were observed in the fertilized eggs of A. zuluensis indicating that sperm with long nuclei might be favoured for fertilization. Spermiogenesis involves: (a) development of the acrosome from a proacrosomal granule; (b) development of the nucleus, characterized by elongation and streamlining with a simultaneous condensation of chromatin; (c) development of the axoneme from the centriole; (d) amalgamation of individual small mitochondria to form elongated mitochondrial derivatives in which cristae are arranged into regularly spaced lamellae; and (f) elimination of cytoplasm. The presence of a manchette, a transient microtubular organelle, which surrounds the acrosome, nucleus and mitochondrial derivatives, is a characteristic feature of spermiogenesis. The gross morphology of the reproductive tract in both male and female cicadas exhibits an organization similar to that in most oviparous insects. The non-functional spermatheca is the only exceptional feature in the female reproductive tract. Its role has been taken over by the common oviduct which, subsequently, has become modified into a swollen, differentiated structure with a dual role of receiving oocytes from the paired ovaries and storage of spermatozoa. Testis mass varies between cicada species; this variation might be linked to the intensity of sperm competition which has been found to be positively correlated with relative investment in spermatogenesis. Based on the preliminary findings of this study, K. punctigera, with its larger testis relative to body size, would be the ideal candidate to show the greatest levels of sperm competition. Accessory glands in both male and female A. zuluensis, D. biconica, P. hirtipennis and O. quadraticollis are very long; this character might be of phylogenetic significance. Despite being notoriously refractory spermiocladistics is potentially valuable in systematic and phylogenic studies of cicadas, especially at the subfamily level.
Identifier Chawanji, A.S. (2007) Spermiogenesis, sperm ultrastructure and reproductive tract morphology in cicadas : implications for systematic relationships. PhD thesis, Rhodes University.