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Die Embryonalentwicklung der Paradiesschnecke ... - TOBIAS-lib

Die Embryonalentwicklung der Paradiesschnecke ... - TOBIAS-lib

Die Embryonalentwicklung der Paradiesschnecke ... - TOBIAS-lib

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Kapitel 2<br />

side of the visceral sac forward and into the mantle cavity, this right-hand<br />

side epithelium in Pt 2+ -exposed snails became the outer epithelium of the<br />

snail’s visceropallium instead of just lining the mantle cavity. The ctenidium<br />

developed on the right-hand side of the visceral sac and should conventionally<br />

also be pushed into the mantle cavity by the growing mantle epithelium.<br />

Instead, in Pt 2+ -exposed M. cornuarietis, it remained at the posterior end<br />

of the snail and even shifted a bit to the left.<br />

All this suggests that in a shell-less and clearly “nonrotated” M. cornuarietis<br />

embryo characteristic traits of both “torted” and “untorted” (in the<br />

sense of the ancient mollusc type) gastropods could be found. Interpretations<br />

are difficult, because the concept of ontogenetic torsion was described<br />

in species with planktonic veliger larvae in which a horizontal rotation is<br />

clearly visible. Nevertheless, based on the facts that the M. cornuarietis visceropallium<br />

also shows a clearly visible horizontal rotation and that torsion<br />

was defined to be a common feature of all gastropods, we presume that the<br />

rotation of the visceral sac in M. cornuarietis corresponds at least to some<br />

extend to the conventional torsion in vetigastropods and patellogastropods.<br />

Our observations lead us to the conclusion hat this torsion-like rotation in<br />

M. cornuarietis is not a unitary process in one fell swoop and only partly a<br />

result of differential growth, as intestine and anus reached their normal positions<br />

without a horizontal anticlockwise rotation of the visceral sac (and,<br />

furthermore, in spite of a vertical rotation), and that this differential growth<br />

affects particularly the outer organs like the ctenidium.<br />

There have been reports on several gastropod species, in which not all components<br />

of the visceropallium rotated synchronously (summarized by Page,<br />

2003); and Page (1997) herself reported a “partial torsion” in Haliotis kamtschatkana<br />

due to a dislocation of pallial tissue which resulted in the mantle<br />

cavity and the anus being located on the right side of the embryo despite<br />

a full 1808 rotation of the shell. She also found this phenomenon in the<br />

gastropods Trichotropis cancellata, Lacuna vincta, Haminoea vesicula, Pleurobranchea<br />

californica, and Diodora aspera (Page, 2003). In T. cancellata,<br />

she also observed full streptoneury of the pleurovisceral connectives despite<br />

a rotation by mere 90 ◦ of the visceral sac. She proposed that torsion should<br />

75

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