Properties of organizer cells change as they migrate from the dorsal lip

Organizer cells begin the process of formation of the dorsal lip of the blastopore. Specifically, these cells are at the center of the dorsal lip and involute, moving forward on the midline (i.e., the middle of the anterior–posterior axis). The first organizer cells to enter the embryo migrate anteriorly to become the head endoderm and head mesoderm. Here they induce neighboring cells to participate in making structures of the head. Organizer cells that involute later in gastrulation induce structures of the trunk, and the last of the organizer cells to move inward from the dorsal lip induce structures of the tail. How do the organizer cells change to enable them to induce head, trunk, or tail structures?

Inductive tissue interactions can suppress as well as activate. Goosecoid is a transcription factor expressed by early organizer cells. Goosecoid activates genes encoding certain signaling factors. As the early organizer cells move forward in the blastocoel, they come into contact with new populations of cells that produce several different growth factors. For head structures to form, certain of these growth factors have to be suppressed. The most anterior organizer cells, under the influence of Goosecoid, produce and release antagonists to those growth factors.

The induction of trunk structures requires suppression of a different set of growth factors. In organizer cells that involute later than the head organizers, Goosecoid is no longer the dominant transcription factor, and these cells express different growth factor antagonists. The induction of tail structures requires still different activities of the organizer cells that involute last. Thus the organizer cells express appropriate sets of growth factor antagonists at the right times to achieve different patterns of differentiation on the anterior–posterior axis.

The initiation of nervous system development also involves a suppressive tissue interaction. For a long time it was thought that the involuting organizer cells actively induced the overlying ectoderm to form nervous tissue rather than becoming epidermis. We now know, however, that epidermis is not the default state of the dorsal ectoderm. Rather, the underlying mesoderm secretes factors called BMP proteins that induce the ectoderm to become epidermis. The role of the involuting organizer cells is to block that induction, allowing the overlying ectodermal cells to follow what is really their default pathway—differentiation into nervous tissue.