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Mechanisms that enable injury responses to prompt regenerative outgrowth are not well understood. Planarians can regenerate essentially any tissue removed by wounding, even after decapitation, due to robust regulation of adult pluripotent stem cells of the neoblast population. Formation of pole signaling centers involving Wnt inhibitors or Wnt ligands promotes head or tail regeneration, respectively, and this process requires the use of neoblasts early after injury. We used expression profiling of purified neoblasts to identify factors needed for anterior pole formation. Using this approach, we identified  zic-1 , a Zic-family transcription factor, as transcriptionally activated in a subpopulation of neoblasts near wound sites early in head regeneration. As head regeneration proceeds, the Wnt inhibitor  notum  becomes expressed in the newly forming anterior pole in  zic-1 -expressing cells descended from neoblasts. Inhibition of  zic-1  by RNAi resulted in a failure to express  notum  at the anterior pole and to regenerate a head, but did not affect tail regeneration. Both injury and canonical Wnt signaling inhibition are required for  zic-1  expression, and double-RNAi experiments suggest  zic-1  inhibits Wnt signaling to allow head regeneration. Analysis of neoblast fate determinants revealed that  zic-1  controls specification of  notum -expressing cells from  foxD -expressing neoblasts to form the anterior pole, which organizes subsequent outgrowth. Specialized differentiation programs may in general underlie injury-dependent formation of tissue organizing centers used for regenerative outgrowth.

zic-1 Expression in Planarian Neoblasts after Injury Controls Anterior Pole Regeneration