The fate of received sperm in the reproductive tract of a hermaphroditic snail and its implications for fertilisation.

Multiple mating, sperm storage and internal fertilisation enhance sperm competition. The great pond snail can use stored sperm for over three months, and frequently mates with different partners. This hermaphrodite, Lymnaea stagnalis, can also self-fertilise and often produces egg masses containing both selfed and outcrossed eggs. Hence, a sperm recipient may exert considerable control over paternity. Using microsatellite markers, we show that when allosperm are present, all genotyped eggs are cross-fertilised. We also find that sperm have the opportunity to compete, because double matings lead on average to equal paternity for each sperm donor. This indicates that received sperm are randomly mixed in storage. To gain further insight into the mechanisms underlying the process of sperm storage, digestion and utilisation, we investigated the fate of donated sperm at different times after copulation. We find that within 3 h after transfer most sperm have been transported into the sperm-digesting organ. Fluorescent labelling of sperm in histological sections further reveals that allosperm are not stored in the fertilisation pouch, but upstream in either the hermaphroditic duct, seminal vesicles, or ovotestis. Besides contributing to the understanding of the mechanisms underlying sperm competition and/or cryptic sperm choice, this study shows that mixed mating cannot be treated as a separate issue in hermaphroditic animals. © Springer Science+Business Media B.V. 2008

Intra-specific morphological variation of the spermatheca in the simultaneously hermaphroditic land snail Helix aperta

In the majority of internally fertilizing animals, females are equipped with sperm storage organs where they store the sperm received during copulation. In many simultaneously hermaphroditic pulmonates, these organs consist of complex spermathecae that show inter- and intra-specific variation in their structure. This variability is theoretically predicted by postcopulatory sexual selection in the context of sperm competition and cryptic female choice. In this study, the variation in the structure of the spermatheca was investigated in the land snail Helix aperta from four natural populations near Bejaia in northern Algeria. The populations were different in local snail density, probably also reflecting the intensity of sperm competition. We tested whether the spermatheca showed differences that are predicted by sperm competition theory. In addition, we tested whether the spermathecal structure depends on the shell size and/or is correlated with other reproductive organs that are thought to be affected by sexual selection. We found that the fertilization pouch of H. aperta consists of a simple fertilization chamber and 3–9 spermathecal tubules. The four populations did not differ significantly in the mean number of these tubules. However, significant differences were found in the length of the main tubule, the length of the fertilization chamber, and the average length of lateral tubules. In addition, strong associations were detected between the lengths of these structures and the local snail density, while no effect of shell size or reproductive organs was found. Our results indicate that the intensity of sperm competition may not affect the total number of spermathecal tubules, but may increase their lengths. This increase in spermathecal length may reflect an improved sperm storage capacity that is probably beneficial in situations of high sperm competitions intensity

Among- and within-population variation in sperm quality in the simultaneously hermaphroditic land snail Arianta arbustorum

Sperm competition models on the evolution of sperm size assume associations with another sperm quality trait, sperm longevity. Sperm length can also provide an indication of possible mechanisms affecting motility and thus fertilization success. Despite their importance, however, detailed mechanisms of sperm competition at the gamete level are poorly understood. In simultaneously hermaphroditic land snails, sperm traits and cryptic female choice are assumed to be crucial in determining fertilization success. We examined the variation in sperm length and number among individuals from four natural populations of the land snail Arianta arbustorum, a species with multiple mating and long-term sperm storage. We also assessed variation in velocity, motility and longevity of sperm in snails from two of the four populations. Independent of shell size, sperm length differed among populations and, to a minor extent, even among individuals within populations. Mean sperm length of a snail was not correlated with the number of sperm delivered in a spermatophore. The mean sperm velocity (=VCL) did not differ between snails from two populations. However, VCL varied among snails. Percentage motility and longevity of sperm differed between snails from the two populations. No correlations were found between length, velocity, percentage motility and longevity of sperm. To conclude, individual snails differed in sperm quality, and this variation may partly explain the differential fertilization success between A. arbustorum snails. Moreover, our findings did not support the positive association between sperm length and longevity assumed by sperm competition models for internally fertilizing species