Properties of the primary organization field in the embryo ofXenopus laevisV. Regulation after removal of the head organizer, in normal early gastrulae and in those already possessing a second implanted organizer

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Abstract
Patterns of individuation occurring in the primary embryonic axis of Xenopus following excision of the organizer region of the early gastrula are described. In some 70% of cases the information for induction of the complete head is qualitatively restored by the time of cell determination, giving rise to an essentially normal embryo. In some 40% of cases a second posterior axis of bilaterality is formed, causing development of a secondary anus, tail-fin and spinal cord, and often somites. The probabilities of twinning in the tailfield and of failure to complete apical regulation (= head formation) are largely independent. After such excision of the head organizer region, a delay of some 3 h in the schedule of visible differentiation in the neurula/tail-bud embryo is commonly incurred, whether or not apical regulation is successful. When the apex is excised from a host embryo which has already contained for some hours a second apex (= head organizer) as described in an earlier paper, that grafted apex then captures a considerably increased territory in the host material, as seen from the size of the individuation field finally caused by it. Such a shift across host cells, of the boundary between fields of positional information due to two organizers, is not seen under any conditions where these are left intact, or where host excision is carried out soon after implanting the donor organizer. In discussing the results and reconciling them with earlier observations, it is shown that they strongly suggest the presence of local polar (i.e. vectorial) properties in the presumptive mesoderm, due to signals from restricted regions which have achieved a special apical state. Repolarization of cells by a new organizer is not very rapid, and may spread decrementally from the source. Data on further delays in development, caused by the presence of the second organizer during regulation in the host apex, suggest that one organizer may act directly on cells elsewhere to delay or prevent the restoration of the apical state there.

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