Developmental

Biology XL
138
Week 2
Pattern
Formation
Zygote  multicellular organism

Kahn
Academy
Embryogenesis sets up the adult body plan

Anterior -----------------------------
Posterior
Questions to be answered

•When and how do the

anterior/posterior and
dorsal/ventral axes become
established?

•How are the segments divided?

•How are the segments specified?

Does each part (segment) develop

individually, or does a body plan
develop in a broad pattern first
and then the details are
refined???
For their discoveries concerning the genetic
control of early embryonic development.
Nobel Screens: Look for lethal mutations
using cuticle as a marker
 Nobel Screens: Look for lethal mutations using
cuticle as a marker:
A/P axis

W mutan W mut W mut W mutan

T t T . T . T t
D/V axis
Drosophila egg has
polarity
 Maternal genes specify the
axes

Maternal effect genes - expressed in mother’s

ovaries and produce mRNAs or proteins that are
deposited into different regions of the egg.
Maternal genes specify the
axes
 Syncytiu
m

A P

bicoi cauda
d l
 Maternal vs. Zygotic
genes

Maternal genes:
Genotype of the mother determines the phenotype of the embryo,
irrespective of the genotype of the father.

A recessive maternal effect mutation will produce mutant animals only

when
the mother is homozygous mutant.

Zygotic genes:
Genotype of the embryo determines its phenotype.
Hierarchy of gene actions: broad pattern first,
details later
Broad
pattern

Finer
details
Three classes
of maternal
effect genes
specify the
anterior-
posterior axis.
 bicoid (TF)
Geneti
is c
necessary
for anterior
structure
formation

Physical
experiments

Experiments demonstrate that the A/P axis is specified by localized cytoplasmic

determinants.
 bicoid mRNA is localized
at the anterior end of
the embryo

bicoid mRNA is localized at anterior.

bicoid protein is expressed in a

gradient.

Injection of purified bicoid mRNA was

sufficient to induce anterior
structures. Driever et al., 1990
Increasing the number of copies of
maternal bicoid results in bigger
anterior domain

Head fold is earliest marker of A/P axis.

A2. wild type at 65% embryo

length. A1. remove one copy of
bicoid.
A3. add one copy of bicoid.
A4. add two copies of
bicoid.

bicoid protein level increased

with increasing
doses of bicoid gene copies.
nanos determines the posterior patterns

nanos mRNA is
localized to the
posterior end of
the embryo.

nanos protein forms

a gradient with the
highest concentration
at the posterior end.
 nanos inhibits the translation of maternal
hunchback mRNA

nanos
The secondary maternal gradients:
maternal hunchback and caudal
The combination of primary and secondary
gradients specify the anterior/posterior axis
Broad
pattern

Finer
details
Zygotic genes in A/P patterning:
Gap genes – first to be
activated by maternal genes
 The gap
mRN
genes A
•Expressed in one to two
domains each by
nuclear cycle 13.

•Divide the embryo into

broad regions

•Mutation leads to the

deletion of several protei
consecutive segments. n

•Encode transcription
factors.
bicoid protein level establishes zygotic hunchback
expression
bicoid is a
morphogen!
 What is a
morphogen?
Alan Turing 1952:
“These factors will be called morphogens, the word being intended
to convey the idea of a form producer”

Text book definition:

A morphogen is any substance active in pattern formation whose
spatial concentration varies and to which cells respond differently at
different threshold concentrations.
hunchback protein gradient helps to
establish other gap gene
expression
knirps mRNA
(black)

normal

hunchback
mutant

hunchback protein can both activate and

repress.
 Gap gene mutual repression
maintains their gradients

hunchback
KrŸppel knirps giant
hunchback

giant

anterior
Broad
pattern

Finer
details
Broad
pattern

Finer
details
dorsal nuclear protein gradient specifies the
drosal/ventral axis

dorsal promotes ventral

fate!!!
The dorsal /ventral axis
is specified by a
ventral nuclear dorsal
gradient
The expression of zygotic genes along the D/V
axis is controlled by Dorsal protein

Dorsal = TF, can act both

as
a repressor and activator
Dorsal target genes have different affinity for the
Dorsal protein

Dorsal represses expression

(dpp)

High affinity + other TFs

(rhomboid)

Low affinity (snail,

twist)