Vertebrate Development

• Organogenesis
– basic body plan established
– tissues develop into organs
– embryo will grow to be a hundred times larger
Vertebrate Development
Vertebrate Development
Gastrulation and Neurulation

• Rearranges the cells of the blastula

• Formation of Germ Layers
• Neurulation involves the process
of forming a neural tube. The tube
is a forerunner of the central
nervous system.
Gastrulation Neurulation
 Organogenesis

• The various organ systems of the embryo are

formed from the three germ layers.
• For vertebrate embryos, one of the earliest
events is formation of the neural tube
(neurulation), which becomes the central axis of
the embryo, around which other development
takes place.
• The major organ systems are formed, all at
roughly the same time.
 Organ formation
• After gastrulation, cells in each of the 3
layers begin to differentiate into tissues
and embryonic organs
Primordia of organs/systems are laid down
– Notochord –becomes spine
– Neural tube –becomes brain and spinal cord
– Somites –give rise to segmented structures
– Coelom –body cavity
 Induction initiates organ formation
• The effect of induction is to switch on
genes whose expression makes the
receiving cells differentiate into a specific
tissue
– Leads to increasingly greater specialization of
cells as organs begin to take shape
 Pattern Formation

• Master control genes respond to chemical

signals that tell a cell where it is relative to
other cells in the embryo
– These signals determine which master control
genes will be expressed
• Ultimately, which body parts will form
How close is the relation between the repeating pattern of
vertebrae & ribs along the human body axis and the body
segments that characterize centipedes, etc ?

This idea of segmentation shows up most obviously

in the organization of the mesoderm into SOMITES
seen along either side of the developing neural tube
Specialization of the Mesoderm
• First evidence is the appearance of the
notochord
• Three mesoderm aggregates appear
lateral to the notochord

– Somites, intermediate mesoderm, and double

sheets of lateral mesoderm
Four kinds of embryonic mesoderm
Axial (notochord)
Paraxial
Forms the axial
skeleton, the
volantary
musculature, and
part of the dermis

Intermediate
Forms the
urinary and
genital systems

Lateral Plate
Somatopleur: body cavity and most of
dermis
Splanchnopleur: Viseral organs
Paraxial, intermediate, and lateral plate
mesoderm.

Axial skel.
Vol muscl.
Dermis

Splanchnopleuric Urinary system

Somatopleuric Genital system
mesoderm
Form ~44 pairs of somites then
caudal most 7 somites disappear
giving rise to 37 pairs.

1-4 somites: occipital part of the

skull, bones of nose and eyes, &
muscles of the tongue.

Next 8 pairs: form in the

presumptive cervical region. Give
rise to occipital bone and cervical
vertebrae, and assoc. muscles.

Next 12 pairs: Thoracic somites …

thoracic vertebrae, and associated
muscles.

5 lumbar somites, 5 sacral somites,

&

Finally 3 coccygeal somites.

 SOMITES AND THEIR FATE
• The 42--44 pairs of Somites have three
functional parts:
– Sclerotome – produce the vertebrae and ribs
– Dermatome – help form the dermis of the skin
on the dorsal part of the body
– Myotome – form the skeletal muscles of the
neck, trunk, and limbs
Specialization of the Mesoderm
• Intermediate mesoderm forms the gonads
and the kidneys
• Lateral mesoderm consists of somatic and
splanchnic mesoderm
Specialization of the Mesoderm
• Somatic mesoderm forms the:
– Dermis of the skin in the ventral region
– Parietal serosa of the ventral body cavity
– Bones, ligaments, and dermis of the limbs
• Splanchnic mesoderm forms:
– The heart and blood vessels
– Most connective tissues of the body system
Neural tube Paraxial mesoderm/somite

Intermediate
mesoderm

Axial mesoderm/Notochord
Somites are not at the far head OPEN
end, because the head gets its
mesenchyme from neural crest 1

The somites are covered with C SOMITES

ectoderm (not shown), with L
O
which part of the somite will S
later interact E
D 7

Remember 7-12-5-5-4
vertebrae. Many more somites
than 7 need to be recruited
N
& some use will be abortive OPE
Subdivisions of the mesoderm
 Neural groove

Somitic mesoderm Regions of mesoderm 1

NC

Endoderm PARAXIAL

INTERMEDIATE

LATERAL
 Regions of mesoderm 2

NC

Clumped as somites PARAXIAL

INTERMEDIATE

Continuous as lateral plate LATERAL

 Roles of mesoderm 1

NC

Paraxial migrates PARAXIAL

around the notochord
INTERMEDIATE
Intermediate caudally makes
tubes to start the kidney LATERAL

Lateral splits for: muscles, skin, & some bone; a deeper part
for the support & construction of visceral organs
 Role of the notochord in specifying somitic cells

A signal from the notochord induces

sclerotome formation

Suppress
dermomyotome, and
induce sclerotome
The fate map of a somite in the chick embryo

(express Pax3)

Migrating
muscle
cells

(express Pax1)

Dermis and
all the trunk
muscle

Cartilage of the
vertebrae and ribs
Somites subdivide into three kinds
of mesodermal primordia.

Dermatome

Myotome

Day 22

Day 28 Day 31
Somites break down
the dorsalateral aspect
flattens and becomes
dermomyotome. The
ventralmedial aspect
undergoes an epithelial
to mesenchymal
transition and becomes
the sclerotome.

Dermomyotome breaks
down into the myotome
and the dermatome
 Molecules that participate in the
specification/differentiation of somites:
Approximately 35 days
 The Process of Neurulation

• Inductive interactions between notochord and

ectoderm
• Ectodermal germ layer differentiates:
– Outer epidermis
– Neural crest
– Neural tube
• Development of the nervous system
 Formation of the Neural Tube

• Primary Neurulation
1. Differentiation of neural plate
2. Shaping of neural plate
3. Bending of neural plate: Neural
folds/Neural Groove
4. Closure of neural groove

• Secondary Neurulation
CAVITATION OF THE MESODERMAL CAUDAL
EMINENCE SECONDARILY JOINING THE NEURAL CANAL
 Formation of the Neural Tube

• Primary and Secondary Neurulation

PRIMARY NEURULATION SECONDARY NEURULATION

Induction of the neural plate.
 Formation of the neural tube

Day 18

Day 19..21

cell proliferation
&
cell specification

Day 22…24
 Neurulation accomplishes three
major things
(1) It creates the neural tube, which gives rise the
central nervous system.
(2) It creates the neural crest, which migrates
away from the dorsal surface of the neural
tube, and gives rise to a diverse set of cell
types.
(3) It creates the bona fide epidermis, which
covers over the neural tube once it is created.
 In the Beginning ….

• Germinal layers:
Skin and Brain
– Ectoderm
Connective Tissues
– Mesoderm
Inner Tubes
– Endoderm
• Neural Ectoderm
– The Neural Plate
• Neural Tube and Neural Crest
Neural Plate (~day 16)
Neural Groove and Folds
Neural Crests
Sinking into the Mesoderm
Neurulation!

Neural Crest
Neural Induction
Neurulation
Neurulation

The neural plate forms after

gastrulation is completed.

The neural tube narrows

along its medial-lateral
Axis. The plate begins to
role into a tube. The
cells at the midline
produce a medial hinge
point (MHP).

As the tube forms and

segregates into the embryo,
neural crest cells emigrate
from the dorsal aspect of
the neural tube.
Day 26
Neural plate

Neural groove

SOMITES
Expansion of the Neural Plate occurs between 18-20 days
Neural Tube formation
Cut edge of amnion

Seen from Dorsal Aspect

Neural tube closes in the middle and zippers closed
Cranially and caudally

Remains opens
to amniotic
cavity
The neural folds fuse,
forming the portion of the
neural tube that will be the
brain and the spinal cord
 Secondary Neurulation occurs caudal to S2 (somite 31)
Caudal eminence

J20 J40

•Gastrulation through the caudal part of primitive streak produces

the mesodermal caudal eminence by day 20
•The caudal eminence cavitates to form a lumen that joins the
neural canal to complete the caudal neural tube by week 8
 Differentiation of Neural Tube
• Major morphological changes: differentiation
of brain ventricles and spinal cord
• Differentiation of neural tube cells
• Development of peripheral nervous system
 Hindbrain (rhombencephalon)

Midbrain Neural Tube

(mes-
encephalon) Notochord

Forebrain (prosencephalon)
 I II III IV V VI
Somitomeres
Somites

Somite development
(mesoderm)
Events in neural crest formation

Ectodermal Thickening

Specialization in the
lateral neural plate

Separation from ectoderm

AND the forming neural
tube

Change of crest cells to a

migratory nature/phenotype

Cells start moving, partly by

their own program, & partly
in response to cues
Closure of the neural folds forms neural tube

Neural crest migrate from edges of neural folds

Neural crest cells
undergo epithelial
to mesenchymal
transition.

Cells delaminate
from sheet and
start to migrate.
 ECTODERM
Derivatives of
GERM LAYERS
MESODERM
ENDODERM
Resulting tubular/ cylindrical form

ECTODERM

MESODERM

ENDODERM