Determination

of Embryonic
axes (II)
Bio 209
PJT Denusta
Biology Department
West Visayas sState University

Left/right axis in Xenopus laevis

Molecular mechanism involves:
Expression of nodal gene Xnr-1
concentration of Xnr-1 dependent on clockwise rotation of
cilia in organizer region
Xnr induces expression of Pitx2 gene

But, what limits Xnr 1 expression solely on the left-hand

side?
The concentration of Nodal protein to the left side is
caused by the clock-wise rotation of cilia found in the
organizer region
This cilia are formed during the later stages of gastrulation
at the dorsal blastopore lip

Although the developing tadpole looks

symmetrical, the internal organs (e.g., heart and
gut) are not evenly balanced
In addition to its dorsal-ventral and anteriorposterior axes, the embryo has a left-right axis.
In all vertebrates studied, the crucial event is the
expression of nodal gene in the lateral plate
mesoderm on the left side of the embryo
in Xenopus, this gene is Xnr1 (Xenopus nodalrelated 1)
If Xnr 1 expression is permitted to occur on the
right-hand side, the position of the heart (normally
found on the left) and the coiling of the gut are
randomized

 Pitx2 determines the direction of heart looping and gut

coiling
If embryo is injected with Pitx2 so that this protein is
present in the mesoderm of both right and left sides,
heart looping and gut coiling is random with respect to
each other
Sometimes treatment results in complete reversals, in
which heart loops to the left and the gut coils in
clockwise manner

Axis determination in the chick

Dorsoventral axis due to differences in pH of
subgerminal space and albumin
Anteroposterior axis traced to the rotation of
the ovum in the hens reproductive tract
creating posterior marginal zone (point where
2ndary hypoblast forms)
Dorsoventral axis of the frog determined by
Primitive streak forms against the posterior side

The Role of gravity in determination

of anteroposterior axis

Lighter yolk

Juncture of area pellucida and area opaca

posterial marginal zone (PMZ)
As the ovum passes through the hens
reproductive tract, it is rotated for about 20 hours
in the shell gland (15 rev/hour)
This shifts the yolk such that its lighter
components lie beneath one side of the
blastoderm.
This imbalance tips up one end of the blastoderm,
and that end becomes the posterior marginal zone
(PMZ), where primitive formation begins
PMZ =Nieuwkoop center

PMZ equivalent to amphibian Nieuwkoop

center
When grafted at anterior region of marginal
zoneinduces primitive streak and H. node but
did not contribute to the cells in both
PMZ produces Wnt8c (capable of inducing B
catenin)
PMZ secrete Vg1 (member of TGF-B family)
PMZ and Vg1 induce Nodal protein
Function of Nieuwkoop center secretes dorsomesodermal inducing factor

Hensens node and derivatives as 1o

organizer
Secrete Chordin, Noggin, and Nodal
proteins
Antagonize BMPs and dorsalize ectoderm
FGFs needed to inhibit effect of BMP4
BMP4 anti-neuralizing molecular

Left and right

axis in Avian
model

Through a gradient of Retinoic acid

lefty1 will be produced all structures that
will be formed here will be the left side
Lefty1 will inactivate hypoblast factor A
Sonic hedgehog protein CARONTE
which will inactivate bmp nodal and
lefty will be expressed pitx2 on all left
side structures will be expressed

Axis determination in Mammals

Anterior-posterior axis- have 2 signalling
centersthe node and the anterior visceral
endoderm (AVE); serve as source of
organizing molecules
Chordin, Noggin from the node
Hesx1, Lim 1, Ot X-2, and Cerberus from AVE

dependent on expression of Hox genes

Dorsal- ventral axis determination

Second polar body supposed to be released in

the area with less yolk
Kung saan ang nucleus which releases the
second polar body is the animal pole and
opposite to the is the vegetal pole

Left and right axis

determination in
Mammals

Ciliary movement in node

Rearrangement of Structures to fit internal

development

biology.kenyon.edu/.../Chap14/Chapter_14.html

End of Topic 7