Cell Biology of the Tardigrades: Current Knowledge and Perspectives.

The invertebrate phylum Tardigrada has received much attention for containing species adapted to the most challenging environmental conditions where an ability to survive complete desiccation or freezing in a cryptobiotic state is necessary for persistence. Although research on tardigrades has a long history, the last decade has seen a dramatic increase in molecular biological ("omics") studies, most of them with the aim to reveal the biochemical mechanisms behind desiccation tolerance of tardigrades. Several other aspects of tardigrade cell biology have been studied, and we review some of them, including karyology, embryology, the role of storage cells, and the question of whether tardigrades are eutelic animals. We also review some of the theories about how anhydrobiotic organisms are able to maintain cell integrity under dry conditions, and our current knowledge on the role of vitrification and DNA protection and repair. Many aspects of tardigrade stress tolerance have relevance for human medicine, and the first transfers of tardigrade stress genes to human cells have now appeared. We expect this field to develop rapidly in the coming years, as more genomic information becomes available. However, many basic cell biological aspects remain to be investigated, such as immunology, cell cycle kinetics, cell metabolism, and culturing of tardigrade cells. Such development will be necessary to allow tardigrades to move from a nonmodel organism position to a true model organism with interesting associations with the current models C. elegans and D. melanogaster.