Organizer effect and anteroposterior patterning

Hensen's node

also known as Primitive knot

influences the regionalization of the ectoderm and mesoderm of the

chick

homologous to the dorsal lip of the blastopore in amphibian embryos

In early stages of development

- the organizer it is not termed as "dorsalization" or "dorsoventral" because
the
chick blastoderm is flat
- the axis is homologous to the axis of Xenopus and zebrafish; and
- runs through the blastoderm (dorsal surface) to the subgerminal cavity
After ventral closure:
- the notochord, somites, and lateral plate run from dorsal to ventral thus, the
nomenclature becomes the same for the chick and the amphibian

Node grafting

Node grafted into the area pellucida (not far from the host streak) can
induce a partial secondary axis
where:
notochord = graft-derived
neural tube and somites = host-derived

resembles the behavior of Xenopus' organizer and shows that the node
emits inductive signal

The activity of the node persists until about H & H stage 8 (four
somites)
where:
younger nodes = induce anterior neural structures
older nodes = induce posterior structures

After H & H stage 4 (definitive streak), the competence of the epiblast

disappears already
Note: Epiblast is able to respond to a graft by generating a complete
embryonic axis where the host epiblast changes its fate but rapidly declines
as soon as the head process begins to emerge.

Preneural markers, Sox3 and Otx2, appear widely expressed at early

stages but does constitute a specification of neuroepithelium as
epiblast explants express Sox2 without signals from the node and do
not progress to actual neuroepithelium

Organizer effect
- due to BMP inhibition
several BMPs are expressed in the peripheral part of the blastoderm
(area opaca)

Chordin - a BMP inhibitor expressed in the node

Reduction of Smad1-phosphate - detectable by immunostaining in

the prospective neural plate region

BMPs can suppress somite formation in favor of the lateral plate as far
as effects on mesoderm are concerned

Noggin - a BMP inhibitor that induces ectopic somites when applied

Note: In general, BMP inhibitors do not have expected neuralizing activity

when applied to explants of epiblast
*Factor: FGFs are responsible for neuralizing activity
FGF inhibitors and Wnts suppress neural induction

Anteroposterior patterning

FGFs are also responsible for the formation of the trunk-tail parts in
anteroposterior pattern, as in Xenopus and zebrafish
- they are expressed in the primitive streak; and

- an FGF-impregnated bead induce posterior neural tube from the

epiblast
- they induce expression of CDX genes that will in turn induce
expression of
posterior Hox genes or HOX6-13

Embryonic regulation

A complete axis can still be produced -> extirpation of Hensen's node

at an early stage

ADMP (antidorsalizing morphogenetic protein) - produces by the node

which has BMP-like activity and is thought to be responsible for
regulation of proportions in Xenopus

The node reforms from the posterior margin of the defect

*requires the presence of the mid-primitive streak

Then, suppresses formation of other nodes in the vicinity