Pattern formation in 8-cell composite embryos of Xenopus laevis.

We have shown in defect experiments that an 8-cell embryo of Xenopus laevis consists of three kinds of cells, that is, animal, vegetal dorsal and vegetal ventral cells, and that cells of different kinds are distinctly different in their developmental capacity. Complete pattern formation occurs in any defect embryo which contains at least two animal, one vegetal dorsal and one vegetal ventral cell. In the present transplantation experiments, we replaced one or two cells of one 8-cell embryo by those of another to obtain 29 series of composite embryos, in which the cell composition of an embryo and/or the dorsoventral orientation of individual cells differed from those of a normal 8-cell embryo. The resulting embryos were examined macroscopically when controls reached stage 26 (tailbud stage) and later. The results showed that both the two animal dorsal cells or one vegetal dorsal cell could be replaced by animal ventral cells or a vegetal ventral cell, respectively, without any detectable effect on pattern formation, irrespective of the ventrodorsal direction of the ventral cells. On the other hand, replacement of an animal ventral or a vegetal ventral cell by an animal dorsal or a vegetal dorsal cell, respectively, made most composite embryos twins. Twins were also formed when a left-handed vegetal dorsal cell was replaced by a right-handed counterpart and vice versa. In these composite embryos, the dorsoventral orientation of the transplanted cell was different from that of a resident dorsal cell or cells of a recipient, and several lines of evidence show that the dorsal cell transplanted in an off-axis orientation is responsible for twin formation. Thus, dorsal cells have the capacity to form dorsal axial structures at later stages and this capacity is localized on the dorsal side, and endows the cells with polarity. On the other hand, ventral cells did not have this capacity or polarity, judging from the fact that their orientation had no effect on pattern formation. One vegetal dorsal or ventral cell could be replaced by an animal dorsal or ventral cell, respectively, without any marked effect. However, replacement of two vegetal cells by animal ones and of one or two animal cells by vegetal ones resulted in deficiency of vegetal cells and oedema and in deficiency of animal cells and incomplete invagination, respectively. Twin formation in composite embryos with animal dorsal cells in place of animal ventral ones is discussed in consideration of findings in recombination experiments by Nieuwkoop.