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Die Embryonalentwicklung der Paradiesschnecke ... - TOBIAS-lib

Die Embryonalentwicklung der Paradiesschnecke ... - TOBIAS-lib

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Kapitel 2<br />

first tended to invaginate and its central portion tentatively evaginated, thus<br />

forming the mantle anlage. However, this anlage stopped growing before<br />

leading to a rotation of the visceral sac and to mantle formation. In ol<strong>der</strong><br />

embryos, it could be observed that the whole complex associated with shell<br />

secretion and mantle formation (consisting of mantle edge, shell gland, and<br />

mantle anlage) did not remain on the lateral side of the embryo but moved<br />

to the ventral side of the visceral sac. This movement may be due to a<br />

redirection of the counter-clockwise rotation program of the visceral sac.<br />

Apparently, the body axis around which differential growth lets the tissues<br />

rotate had changed (Fig. 10) and caused the whole visceral sac to rotate vertically<br />

by 90 ◦ . The described movement alteration is also in accordance with<br />

an observation we made during our investigations of the internal anatomy of<br />

adult shell-less snails (not yet published): the osphradium, which originates<br />

on the right dorso-lateral side of the visceral sac, was finally located on the<br />

left ventro-lateral side of the visceral sac in Pt 2+ - exposed M. cornuarietis<br />

(unpublished data). This is not a surprise if its anlage rotated vertically by<br />

90 ◦ relative to its origin. The orientation of this altered rotation also explains<br />

why the ctenidium that differentiates next to the osphradium on the<br />

right dorso-lateral side shifted sinistrad relative to its origin.<br />

As Osterauer et al. (2010b) described, the shell gland secreted calcium<br />

carbonate despite its abnormal position. This calcium carbonate precipitation,<br />

called “internal shell”, was found to cup the digestive gland inside the<br />

snail’s body. How these internal shells are secreted exactly is unknown. We<br />

also do not yet know about the future of the mantle anlage during postembryonic<br />

development. With time, the shell grows larger inside the snail, but<br />

shell gland and mantle edge do not expand much. Normally, it would not be<br />

possible for the shell to grow into the body because of the un<strong>der</strong>lying tissue<br />

(mantle anlage). So for this to happen, the invaginated mantle anlage either<br />

followed the growth of the internal shell or it was torn by it as a result of<br />

shell growth, two possibilities one may speculate upon. Demian and Yousif<br />

(1973c) reported that the mantle first appeared in Stage VII, when the rotation<br />

of the visceral sac had begun, as a result of the excessive growth of the<br />

mantle anlage. Lacking this outgrowth, Pt 2+ -exposed embryos neither de-<br />

73

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