PROPORTION REGULATION

o If a material is removed from the embryo, the pattern that eventually forms
does not have a gap but rather consists of a reduced-scale version of the
normal pattern.
o Pattern is usually not perfect but certainly can tend toward the normal
o Exhibits antiregulative behavior due to the accumulating morphogen at the
cut
o Key is the expression of ADMP
o Removal of molecular components that regulate proportion such as BMP
2,4,7 and ADMP leads to total neutralization

ADMP- antidorsalizing morphogenetic protein; a bone morphogenetic (BMP) type

inducing factor

- expressed in the organizer, yet has ventralizing properties

- BMP signaling represses its transcription

ANTEROPOSTERIOR PATTERNING

o Neural-inducing activity is shown both by the organizer and by the axial

mesoderm into which the organizer develops.
o Anterior organizer or anterior axial mesoderm induces brain structures
o Posterior organizer or posterior axial mesoderm induces both brain and
spinal cord
o Posterior signal controls anteroposterior patterning during the formation of
the central nervous system (CNS)
o The two domains within the organizer in the initial anteroposterior pattern
are derived from the different ratio of β-catenin arising from cortical rotation
and nodal-related signaling arising from mesoderm induction

Anterior part- will become anterior ectoderm and prechordal mesoderm

*goosecoid will induce expression of anterior-type genes from ectoderm, such as otx2
(fore/ midbrain) and ag1 (cement gland)

Posterior part- will later become the notochord and somites and, during
gastrulation, it elongates considerably by convergent extension.

*homeobox gene not and the T-box gene brachyury will induce expression of both anterior-
and posterior-type genes from the ectoderm (e.g. both otx2 and Hox genes), and its
posteriorizing activity due to secretion of FGFs and Wnts. Together these upregulate a
group of homeodomain transcription factors encoded by the cdx genes and these in turn
upregulate the posterior Hox genes of paralog groups 6–13.
Evidence that FGF and Wnt signaling is required to induce the trunk–tail region:

1. If animal caps are treated with the BMP inhib- itor noggin, then only anterior-type
neural genes are induced, but addition of Wnt or FGF will also induce posterior
neural genes
2. Overexpression in embryos of a dominant negative FGF receptor that inhibits
endogenous FGF signaling, or of the dick- kopf Wnt inhibitor, will prevent formation
of the posterior.
3. Overexpression or inhibition of retinoic acid does have effects on the pattern but
in Xenopus largely confined to the hindbrain

FGF- effects mostly felt on the trunk-tail region

Wnt- effects reach on the hindbrain region; controls engrailed [midbrain-hindbrain

region]

ORGANIZER GRAFT

o All three of the processes, dorsalization, neural induction, and

anteroposterior patterning are shown here
o A piece of tissue from above the dorsal blastopore lip is implanted into the
ventral marginal zone leading to the formation of a double dorsal embryo.
o The notochord of the ectoderm above the graft becomes induced to form a
second neural tube. As gastrulation proceeds, both host and graft axes form
progressively more posterior parts.
o The final result is a symmetrical pair of embryos joined belly to belly.
o The movements of the grafted organizer are autonomous and
preprogrammed.
o The secondary axes arising from organizer grafts often lack a head, but if the
graft includes the deep anterior region of the organizer then this will emit
cerberus and dickkopf and induce a head in the overlying ectoderm.