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World Congress of Malacology Antwerp ... - Unitas Malacologica

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Gastropod species as model organisms in ecotoxicology ?<br />

Introducing the molecular view<br />

Dallinger, Reinhard<br />

Institute <strong>of</strong> Zoology, University <strong>of</strong> Innsbruck, Austria,<br />

Email: reinhard.dallinger@uibk.ac.at<br />

During their evolutionary history, gastropods have been successful in colonizing different<br />

environments with some <strong>of</strong> the most adverse habitats on the earth. One <strong>of</strong> the preconditions for this<br />

adaptational capacity <strong>of</strong> gastropods has been their huge physiological plasticity, along with their high<br />

ability to cope with stressful conditions <strong>of</strong> different origin. In particular, this remarkable potential for<br />

adaptation is based, among other features, on an energy metabolism with a variety <strong>of</strong> anaerobic<br />

pathways. This enables some gastropod species to survive in habitats with limited energy resources,<br />

and to outlast periods <strong>of</strong> adverse environmental conditions by reducing their activity and energy<br />

demand to a minimum. In addition, a range <strong>of</strong> stress compensation and detoxification mechanisms<br />

ensures the organism’s survival even under the influence <strong>of</strong> stressful physical and chemical<br />

environmental factors, including adverse human impact. All these remarkable features make<br />

gastropods potential candidates for fundamental and applied research in ecotoxicology. In the present<br />

talk, I would like to substantiate this hypothesis by focusing on the exceptional ability <strong>of</strong> some<br />

pulmonate species to cope with metal stress, and perhaps with more than this.<br />

Some terrestrial pulmonates (e.g. Helix pomatia, Cornu aspersum, Arianta arbustorum) possess<br />

mechanisms <strong>of</strong> metal detoxification, which are much more specific compared to those found in most<br />

other animal species. The primary reason for this is the fact that these pulmonates rely on metal<br />

binding Metallothioneins (MT) consisting <strong>of</strong> metal- and function-specific is<strong>of</strong>orms. In the Roman<br />

snail (Helix pomatia), for example, one MT is<strong>of</strong>orm is devoted to Cu homeostasis, while a second<br />

is<strong>of</strong>orm deals with detoxification <strong>of</strong> Cd. The adaptation <strong>of</strong> the two peptides to their metal-related<br />

tasks represents an unique example <strong>of</strong> structural and functional diversification <strong>of</strong> different is<strong>of</strong>orms<br />

within a protein family. Because <strong>of</strong> its metal-specific features, the Cd-MT is<strong>of</strong>orm <strong>of</strong> Helix pomatia<br />

and some other related pulmonate snails has successfully been used as a biomarker <strong>of</strong> exposure in<br />

metal-contaminated terrestrial environments.<br />

Another unique feature <strong>of</strong> the MT system <strong>of</strong> terrestrial pulmonates is the complexity <strong>of</strong> their MT<br />

genes. The Cd-MT gene <strong>of</strong> Helix pomatia, for example, is by far the largest and most complex MT<br />

gene known throughout all organismic kingdoms. Its nucleotide structure suggests that at the<br />

molecular level, adaptation <strong>of</strong> terrestrial pulmonates to metal stress is achieved by a broad potential<br />

<strong>of</strong> response patterns to a diversity <strong>of</strong> extrinsic and intrinsic stress stimuli, rather than strict metal<br />

specificity. This is confirmed by the fact that the Cd-MT is<strong>of</strong>orm gene <strong>of</strong> Helix pomatia contains,<br />

apart from Metal-Responsive Elements (MREs) and nuclear activators and silencers, a manifold array<br />

<strong>of</strong> binding sites for various stress-related transcription factors. These include, among others,<br />

transcription factors involved in heat shock response, xenobiotic metabolism, and immune defense.<br />

The multiplicity and diversity <strong>of</strong> these transcription factor binding sites probably allows, apart from<br />

metal responsiveness, fine-tuning <strong>of</strong> transcriptional activation <strong>of</strong> the Cd-MT gene in dependence <strong>of</strong><br />

many other evnironmental stressors. In fact, the induction rate <strong>of</strong> the Cd-MT gene <strong>of</strong> Helix pomatia<br />

due to metal exposure is much higher at the transcriptional level than at the protein level, a feature<br />

which has also been observed for other stress-related proteins. The high complexity <strong>of</strong> the Cd-MT<br />

gene <strong>of</strong> Helix pomatia and its sensitivity to transcriptional induction and regulation suggests that it<br />

could be used as a complex biomarker for stress response, which includes, apart from metals, many<br />

other environmental stressors. We are currently testing this hypothesis by transfection <strong>of</strong> reporter<br />

gene constructs from Helix pomatia into embryonic cells <strong>of</strong> the freshwater pulmonate Biomphalaria<br />

glabrata.<br />

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