Pattern regulation and the origin of extra parts following axial misalignments in the urodele limb bud.

Pattern regulation following axial misalignments in the stage-38+to stage-40 urodele limb bud was studied on one newt and two salamander species. Grafts of the distal tip of the limb bud were made to the stump of a host limb bud from which a similar piece had been removed. The grafts were positioned with either their anteroposterior, dorsoventral, or both of these axes reversed with respect to the host axes. Mirror-imaged duplications, positioned posteriorly or both anteriorly and posteriorly, occurred nearly all (96%) of the time when the anteroposterior axis was reversed. Dorsoventral axial misalignment rarely promoted the generation of mirror-imaged duplication (8%) but did affect the organization along the anteroposterior axis by causing a serial repetition of either digit 2 or digit 3. Regulation, therefore, does not always occur along each axis independently of the others. Consistent with the data derived from reversing individual axes, most of the duplications which occurred when both axes were reversed were in the anteroposterior plane. Some were in the dorsoventral plane, and a few had intermediate positions. Of these duplications a few were neither right not left hands, rather they were of mixed handedness with a change in the dorsoventral polarity from the anterior border to the posterior border. Whether extra parts which result from axial misalignments arise from the graft, the host, or both the graft and the host was investigated using heteroplastic grafts and grafts exchanged between triploid and diploid axolotls. Duplications were found to have cellular contributions from both the graft and the host. In some cases on source would dominate but usually both made a substantial contribution. The diploid-triploid material suggests that a considerable mixing of host and graft cells may occur in duplications. Additionally, some digits of the graft sequence of digits can be derived from host tissue. The extra digit in those hands displaying a serial repetition was derived from host tissue in some cases and graft tissue in other cases.