Tissue transplantation in planarians: A useful tool for molecular analysis of pattern formation

Freshwater planarians are well known for their remarkable plasticity and regenerative capabilities. Most studies of planarian regeneration have specifically examined regeneration after transverse or longitudinal sectioning or during homeostasis in intact adults. However, tissue transplantation, first performed over a century ago, constitutes another important tool in the study of regeneration in planarians, and can be easily performed given this species' extraordinary healing capacity and its lack of a circulatory system. Studies conducted to date have demonstrated the viability of transplantations involving a variety of tissue types of different positional identities, affecting any of the 3 main body axes. Moreover, these grafting experiments have shown that tissues possess axial positional identities, which are retained following transplantation. The confrontation between different positional identities that occurs after any type of tissue transplantation is resolved by the formation of a blastema, consisting of undifferentiated tissue produced by adult pluripotent stem cells (neoblasts). This blastema intercalates the positional identities of the graft and host tissues. The recent discovery of pathways involved in planarian growth, patterning, and organogenesis, as well as corresponding molecular markers, makes tissue transplantation a vital new tool with which to explore pattern formation. Here, we discuss the different grafting approaches used in planarians, and the corresponding intercalary regenerative response, placing particular emphasis on the respective contributions of donor and host tissue. Moreover, we discuss the temporal induction of blastema formation, and present new molecular data on the generation of an ectopic anterior/posterior axis in response to dorsal/ventral confrontations between host and donor tissue.