Wnt signaling in Pristionchus pacificus gonadal arm extension and the evolution of organ shape.

Changes in organ morphology have been essential to the evolution of novel body forms and in permitting organisms to invade new ecological niches. Changes in the arrangement of cells and tissues and in the regulation of morphological movements are fundamental to evolutionary transitions of organ shape and function. However, little is known about the genetic and developmental control of these changes. We use interspecific differences in the migration and extension of the nematode hermaphrodite gonadal arms to study the generation of morphological novelty. We show that the extending Pristionchus pacificus gonadal arms display a ventral migration that is unique to the Diplogastridae in comparison to the Rhabditidae, including Caenorhabditis elegans, and other nematodes. This results in the distal gonad residing along the ventral side of the body in P. pacificus in contrast to lying on the dorsal side of the body as in C. elegans. We show that at the cellular level this morphogenetic movement is regulated by signals from the developing vulva and the sister gonadal arm. We further show that in P. pacificus Wnt signaling is essential for this regulation. We show genetic and molecular evidence that suggest the Wnt ligands Ppa-mom-2 and Ppa-cwn-2 are components of the signaling mechanism. Supporting these findings, the hermaphrodite gonad of Ppa-bar-1 mutant animals mimics the shape of the C. elegans hermaphrodite gonad; the arms fail to extend ventrally. Thus, this genetic analysis of gonad migration provides insight into the mechanisms underlying the generation of morphological novelty and organ shape.