DEVELOPMENT

   OF    THE  

VERTEBRATE    BODY    PLAN


Thomas  A.  Marino,  Ph.D.  

Temple  University  School  of  Medicine
 DEVELOPMENT    OF    THE  

VERTEBRATE    BODY    PLAN

Early  Development  
1.  Development  of  Ectoderm  
         A.  Neural  Tube  
         B.  Surface  Ectoderm  
2.  Development  of  Endoderm  
         A.  G.I.  Tract  
   B.  Respiratory  Tree  
         C.  Pharynx  
3.  Development  of  Mesoderm  
         A.  Paraxial  
         B.  Intermediate  
 Summary  of  Week  2
Trophoblast Cytotrophoblast SyncyQotrophoblast
Embryonic  Ectoderm
Embryonic    
Mesoderm
Embryonic    
Epiblast Mesoderm
Extraembryonic  
Blastocyst mesoderm
PrimiQve  
Streak Notochordal  
Epiblast Process
Notochord
Embryonic  
Endoderm
Amnioblasts
Inner  Cell    
Mass

Hypoblast Extraembryonic   Yolk  Sac    

endoderm Endoderm
 Trilaminar  Embryo

So by day 18 of gestation,
gastrulation is nearing
completion and the trilaminar Ectoderm mesoderm
embryo will begin:
1.Neurulation
2.Lateral body folding
3.Head and tail folding
!
The trilaminar embryo has three
layers:
Notochord
1.Ectoderm
2.Mesoderm
3.Endoderm Endoderm
 Trilaminar  Embryo

As the third week of gestation is

coming to an end, the mesoderm layer paraxial  mmesoderm
paraxial   esoderm
begins to subdivide into three masses:
1.Paraxial mesoderm
1. This will become the somites.
2.Intermediate mesoderm.
1. This will give rise to much of
the urogenital system.
2.Lateral plate mesoderm.
1. This will become
1. Lateral plate splanchnic
mesoderm intermediate  mesoderm
2. Lateral plate somatic
lateral  mesoderm
mesoderm

 Ectoderm
Neurulation begins with the
notochord cells migrating
around the primitive node.

Notochord

Neural    
Plate
 Ectoderm
As the notochord cells
migrate in a cephalic
direction, the overlying
ectoderm cells will begin Notochord
to differentiate into
neural ectoderm or the
neural plate.
Neural    
Plate
 Ectoderm
Oral  
Plate

The notochord and the neural Prechordal  

plate cells will grow cephalically. Plate
1.This growth will stop in the
region of the prechordal plate.
2.Cephalic to the prechordal Notochord
plate is the oral plate.
1. The oral plate is
Neural    
ectoderm and
endoderm without Plate
intervening mesoderm.
2. The oral plate
demarcates the
separation between the
prospective oral cavity
and the developing gut
tube.
 Neurula'on  –  how  does  
neural  ectoderm  develop?

!
•BMP  gradients  lead  to  formaQon  
of  intermediate  and  lateral  
mesoderm.  
Chordin
Noggin
! Follistatin
•Chordin,  Noggin  and  FollistaQn   !
form  in    cranial  paraxial   !
mesoderm.  They  bind  to  and   !
inacQvate  BMP.   FGF  
! WNT3a
•They  are  present  in  notochord  
and  paraxial  mesoderm  and  
induce  ectoderm  to  become  
neural  plate.  
!
•WNT3a  and  FGF  do  this  in  the  
caudal  paraxial  mesoderm.  
!
•Without  BMP  inacQvaQon  the  
epidermal  phenotype  is  
Neural  crest  forma'on

Neural  crest  cells  form  in  response  to    

! Chordin
! Noggin
Intermediate  levels  of  BMP   Follistatin
! !
!
! BMP !
This  leads  to  PAX3  expression     FGF  
+   WNT3a
FOXD3  expression  is  necessary  for  
neural  crest  formaQon  
And  
SLUG  expression  allows  for  to  neural  
crest  migraQon
Bone  morphogene'c  protein  signalling  and  
vertebrate  nervous  system  development

Aimin  Liu  &  Lee  A.  Niswander

Nature  Reviews  Neuroscience  6,  945-­‐954  
(December  2005)

The  CNS  arises  from  a  specialized  

epithelium,  the  neural  plate  (1).  This  
process  relies  on  the  inhibiQon  of  bone  
morphogeneQc  protein  (BMP)  
signaling.  Folding  of  the  neural  plate  to  
produce  the  neural  groove  is  triggered  
by  the  formaQon  of  a  disQnct  hinge  
point  in  the  ventral  region  (the  floor  
plate;  2).  At  the  end  of  neurulaQon,  
the  lateral  edges  of  the  neural  plate  
fuse  (3)  and  segregate  from  the  non-­‐
neural  epithelium  to  form  a  neural  
tube  (4).  The  roof  plate  and  floor  plate  
form  at  the  dorsal  and  ventral  midline  
of  the  neural  tube,  respecQvely.  The  
roof  plate  becomes  a  new  organizing  
centre  that  produces  BMPs,  which  
provide  dorsal  paaerning  informaQon.  
Neural  crest  cells  derive  from  the  
dorsal  neural  tube  and  migrate  out  to  
form  the  PNS,  as  well  as  melanocytes  
and  carQlage  in  the  head.  Neural  crest  
cells  have  been  shown  to  form  at  an  
intermediate  level  of  BMP  signaling.
 Ectoderm
The  next  stage  of  
neurulaQon  in  the   Neural    
Groove
formaQon  of  the  
neural  tube.    The   Neural    
Plate
lateral  edges  of  
the  neural  plate  
will  form  neural  
folds  and  a  
midline  neural  
groove  appears.
 Fig. 1 (A) Successive images showing the progression of neural tube closure in a stylized
vertebrate embryo (rostral = up).

J B Wallingford et al. Science 2013;339:1222002

Published by AAAS
 Fig. 3 Multiple cell behaviors contribute to neural tube morphogenesis.
Multiple cell behaviors contribute to neural tube morphogenesis. In this schematic, pink and
green cells illustrate convergent extension. By exchanging neighbors specifically in the
mediolateral (horizontal) axis, the sheet of cells is elongated in the anteroposterior (vertical)
axis. Blue cells illustrate apical constriction. These cells do not move but rather change their
shape, leading to a bend in the tissue sheet. [Schematic adapted from (7)]

J B Wallingford et al. Science 2013;339:1222002

Published by AAAS
 Ectoderm

As  the  third  
week  comes  to   Neural Groove
an  end  the   Neural Fold
Somites
neural  folds  
become  
prominent  and  
they  approach  
one  another  in  
the  cervical  
region. Day 20
 Ectoderm
The    neural  tube  is   Pericardial    
first  formed  in  the   Bulge

cervical  region.    It  

then  conQnues  to   Somites
form  in  a  cephalic  
and  caudal  
direcQon.  

Neural  Tube
 Ectoderm
The  last  two  areas  
to  fuse  are  the   Cranial  Neuropore

cranial  neuropore   Pericardial  Bulge

cephalically  and  the  
caudal  neuropore  
caudally.   Somites

Caudal  Neuropore
 UNSW  Embryology

This  link  takes  you  to  the  University  of  New  
South  Wales  embryology  website  where  you  can  
see  an  image  of  an  embryo  at  22  days  of  
gestaQon.    NeurulaQon  is  nearly  complete.    
 Lateral  Body  Folding
• As  neurulaQon  occurs  the  embryo  begins  
lateral  body  folding.  
 Lateral  Body  Folding
• Lateral  body  
folding  involves:  
– The  ectoderm  at  
either  end  
meeQng  in  the  
ventral  midline.  
– The  endoderm  
forming  the  gut  
tube.
 Lateral  Body  Folding
• The  lateral  plate  mesoderm  will  split  into  
somaQc  and  splanchnic  mesoderm.  
• In  between  the  two  the  body  cavity  will  form.  
• This  space  on  both  sides  will  also  fuse  in  the  
midline. Body cavity

Somatic mesoderm
Splanchnic mesoderm
 Lateral  Body  Folding
• As  the  space  between  the  lateral  plate  somaQc  
and  splanchnic  mesoderm  enlarges  the  lateral  
body  folding  conQnues.

Body cavity
23
 Lateral  Body  Folding

Body cavity

Amniotic Cavity

Gut Tube

• The  amnioQc  cavity  will  also  fold  around  the  

embryo.
 Lateral  Body  Folding
• The  two  ends  of  the  ectoderm,  lateral  plate  
somaQc  mesoderm,  lateral  plate  splanchnic  
mesoderm  and  the  endoderm  conQnue  to  
migrate  ventrally.
 Lateral  Body  Folding
• The  amnioQc  cavity  and  the  body  cavity  also  
migrate  ventrally.
 Lateral  Body  Folding
• Now the amniotic cavity surround the embryo which is surrounded by
ectoderm.
• The fused body cavities are surrounded by lateral plate somatic mesoderm
deep to the ectoderm
• The lateral plate splanchnic mesoderm surround the gut tube which is a fused
tube of endoderm

Somatic mesoderm
Splanchnic mesoderm

Body Gut Tube

cavity
 Lateral  Body  Folding
Paraxial  mesoderm Intemediate  
Surface  ectoderm mesoderm
Neural  Tube

AmnioQc  
Cavity

Body Gut Tube

Cavity
Chorionic
Cavity

The embryo is surrounded by the chorionic cavity which surrounds the amniotic cavity.
 Lateral  Body  Folding


This  is  a  very  nice  animaQon  of  lateral  body  

folding  that  will  give  you  a  beaer  three  
dimensional  sense  of  the  process.    However  
this  require  the  flash  plugin.    
 UNSW  Embryology


• This  is  an  image  from  the  UNSW  embryology  

website.    It  shows  a  23  –  26  day  old  embryo  
(week  6  LMP).    The  neural  tube  is  fusing  with  
embryo  is  undergoing  lateral  body  folding.  
• Another  image  can  be  see  by  clicking  here.
 Surface  Ectoderm Neural  
epidermis, hair, nails, Crest:
cutaneous and mammary
glands, anterior pituitary gland, cranial and sensory ganglia
enamel of teeth, inner ear, and and nerves,
lens medulla of adrenal gland,
pigment cells,
branchial arch cartilages,
head mesenchyme

mesoderm
Ectoderm
Neural  Tube

central nervous system,

retina,
pineal body,
posterior pituitary

Endoderm

Ectoderm then develops into three main categories: surface ectoderm, neural crest
derivatives and neural tube derivatives.
 Endoderm
• ConQnuing  the  discussion  of  lateral  body  
folding  and  adding  head  and  tail  folding    The  
development  of  the  endoderm  can  be  
considered  as  it  develops  into:  
– Pharynx  
– GI  tract  
– Respiratory  system  
– Caudal  urogenital  system  structures
 Endoderm  
Ager:  
– Lateral  Body  Folding  
– Head  and  Tail  Folding  
The  endoderm  consists  of:  
– Foregut  
– Midgut  
– Hindgut
 Endoderm
If a sagittal section of the embryo is made and looked at from the side one can see
the ectoderm (blue), mesoderm (red) and endoderm (yellow).
 Endoderm
• Dorsal to the ectoderm is the amniotic cavity and ventral to the endoderm is
the yolk sac.
• The chorionic cavity surrounds these structures.
• The connecting stalk connects the embryo to the placenta.
• The cephalic end of the embryo is defined by the oral plate.

Connecting stalk
Oral plate

Amniotic
Cavity

Yolk
Sac

Chorionic
Cavity
 Endoderm
• The cephalic end of the embryo grows the fastest and as it does the brain moves
into a cephalic position and move the oral plate ventrally and caudally.
• This leaves two cul-de-sacs at either end of the gut tube called the foregut and
hindgut.

foregut
hindgut

ra in
b

Oral plate
 Endoderm
The gut tube begins at the oral plate, starts as the foregut, continues as the midgut
which is still continuous with the yolk sac and then becomes the hindgut ending as
the cloacal plate.
foregut hindgut

midgut

Cloacal  
Plate

Oral  Plate

Allantois
Heart  

Yolk  Sac
 ENDODERM
Cloaca Allantois
A diverticulum from the
ity
connecting stalk called the
  cav
nio
' c allantois connects to the
Am
hindgut and this fusion region
is now called the cloaca.
Uterine
Cavity

Chorionic    
Yolk  
Cavity Sac
The gut tube is divided into three regions:
1.Foregut (yellow) Midgut
2.Midgut

Back
1. Cephalic end (green) Hindgut
2. Caudal end (blue)
3.Hindgut (purple)
1. Cloaca

Tail
Head

Connecting Stalk
Cloaca
Allantois
Foregut
Vitelline Stalk
 Endoderm The endoderm gives rise to four
major components:
mesoderm 1.Pharynx
Ectoderm 2.Respiratory epithelium
3.GI tract and associated glands.
4.Caudal urogenital system

G.I.:  
epithelium of G.I. tract
liver, pancreas
Endoderm !
Caudal  UG  system:
urinary bladder
Pharynx:   Respiratory:   urethra
epithelial parts of: epithelial parts of:
pharynx trachea
thyroid bronchi
tympanic cavity lungs
tonsils, parathyroids
 GastrulaQon
Ectoderm

Amnio'c  Cavity
Paraxial  mesoderm  
!
Intermediate  mesoderm  
!
Lateral  plate  mesoderm

The mesoderm initially Yolk  Sac

is subdivided into three
Endoderm
masses:
1. paraxial mesoderm Notochord
2. intermediate mesoderm
3. lateral plate mesoderm
 Mesoderm
• The  first  part  of  
the  mesoderm  to  
examine  is  the  
lateral  plate   !
!
mesoderm  and   Lateral  plate  
specifically  the    mesoderm:  
Splanchnic   SomaQc  
!
mesoderm !
Splanchnic

42
 Mesoderm
Looking down on the
embryo, the lateral Oral  plate
plate splanchnic
mesoderm begins to Blood    
Islands
see a proliferation of
cells that form blood
islands.
!
These blood island
are forming in the
lateral mesoderm and
continue cephalically
located cephalic to
the oral plate.
 Blood  Islands  
!
!
!
Endocardial    
Heart  Tube

The blood islands consist of angiogenic cell clusters that

will become cells that will form the:
1. endothelium of blood vessels.
2. heart tube.
3. blood cells.
 Mesoderm
Three  sites  of  
early  blood   Chorion

island   Connecting stalk

formaQon:   Cardiogenic area

– cardiogenic  
area  
– yolk  sac  
– chorion  and  
yolk  sac
connecQng  
stalk
 Mesoderm Endocardial
heart tube

Take a cross section in the plane of the dotted line, and looking
at the section from the caudal region, shows the location of the
endocardial heart tubes in the splanchnic mesoderm.
 Mesoderm
The following images
Endocardial
demonstrate that during heart tube
lateral body folding the
endocardial heart tubes will
come together to form the
heart tube.
 Dorsal Aortae

Endocardial
heart tube

The endocardial heart

tubes will come toward
the midline.
!
They will be continuous
with newly forming
dorsal aortae.
By day 21the ends of the
heart tube have fused and
form the heart tube. It is
located ventral to the gut
tube. The first region to fuse
is the prospective ventricular
region.
!
At this point the heart starts to
beat.
 Mesoderm
The fused heart
tube consists of Foregut Dorsal  Aorta
the endocardium
(red) the,
myocardium from
primary and
secondary heart
fields (green
Heart
stipple) and
intervening
cardiac Body  
mesenchyme Cavity
(jelly) in green.

AmnioQc  Cavity
 !
• UNSW  Embryology - This is a scanning electron
microscope image of the fused heart tube.
!
• UNSW Embryology 1 - This is a series of images
showing the fusion of the heart tube.
 Mesoderm
Ectoderm

Paraxial  mesoderm   Amnio'c  Cavity

!
Intermediate  mesoderm  
!
Lateral  plate  mesoderm

The medial mass of

mesoderm is called the
paraxial mesoderm and Yolk  Sac
later the somite.

Notochord
Endoderm
 Dermatome
Myotome

Paraxial mesoderm ->Somite

Sclerotome
The somite will become subdivided into three components:
1. Dermatome - the connective tissue of the skin.
2. Myotome - skeletal muscle cells.
3. Sclerotome - give rise to cartilage and bone of ribs and
vertebrae
 me
y oto
m
rmo WNT
De

AX3
P

Scleretome SHH
(PAX1)

Initially the dermatome and myotome are fused with the

dermatome in between the two masses of myotome. Signals
from the notochord and neural tube influence the
developmental fate of the the somite.
 The medial mass of myotome cells will become the muscles of the back.
The lateral mass of myotome cells will become the muscle of the body wall and
extremities.

Body wall and

Extremity Muscles Back (epaxial) muscles
DermisMYF5

MYOD
 Mesoderm
• Intermediate  
mesoderm  will  
Intermediate  
develop  into   mesoderm   Amnio'c  Cavity
components  of  
the  urinary  
and  
reproducQve  
systems.
Yolk  Sac

56
Development of the

Kidneys
• By day 23 intermediate
mesoderm (orange and
navy) is identified lateral
to the paraxial mesoderm
(red).
• Intermediate mesoderm
is organized into:
• pronephros
• mesonephrose
• metanephros
 Mesoderm
• The  intermediate   Kidney tubules

mesoderm  gives  rise  to:  

– Kidney  tubules  (blue)  
– Ducts  of  the  urinary  and   Primordial germ
cells
reproducQve  systems   Ducts
Body epithelium
– Gonads  
– The  gonads  form  from  
epithelium  covering  the  
intermediate  mesoderm  
(green)  and  migraQng  
primordial  germ  cells. 58
 1. Skeletal muscles,
Urogenital system Mesoderm 2. Skeleton (except skull)
3. Dermis of skin
including :
Paraxial  Mesoderm 4. Connective tissue
1. Kidneys
2. Gonads,
3. Ducts,
4. Accessory glands Lateral  Mesoderm

Intermediate  Mesoderm
1. Connective tissue of
viscera.
2. Serous membranes of:
A. pleura,
B. pericardium
C. peritoneum
3. Blood and lymph cells
4. Cardiovascular system
5. Lymphatic system
• Early  Embryology:  Where  are  we  now  and  
where  are  we  going.  
!

• View the CNN Report

60
 Timing of pregnancy
Embryology/
Gestational Age Clinical Age
Beginning of last Ovarian follicle Day 0
menstrual period matures

Day 0 Proliferative phase Ovulation Day 14

of menstrual cycle
Day 1 Secretory phase of Fertilization Day 15
menstrual cycle.
Day 6 - 7 Implantation Blastocyst Day 20 - 21

Day 14 Primary villi in the Bilaminar disk Day 28

placenta
Day 15 First menstrual Gastrulation Begins Day 29
period missed
 Pregnancy loss
• Approximately 30% of the fertilized eggs
are carried successfully.
• Of the 70% that are unsuccessful almost 1/3
are lost prior to implantation.
• About 40% of postimplantation pregnancies
abort spontaneously,
• Clinically only 10 - 15% are observed.
 Pregnancy loss
• Studies on aborted material demonstrates 50 -
60% have chromosomal anomalies.
• Very early losses closer to 70%
• Higher spontaneous loss in older women.
• Other reasons for loss:
– Genital tract abnormalities.
– Infections
– Endocrine and metabolic anomalies
– Hematologic and immune disorders
 Chances of Conception*

In one month In six months In one year

Early 20's 25% 75% 94%

Late20's/early30's 15% 38-47% 70-85%

Late30's 10% 22-24% 65-70%

* from iVillageHealth.com.
 Average Time to Conception*

No. of months

Early 20's 4-5

Late 20's 5-7

Early 30's 7-10

Late 30's 10-12

* from iVillageHealth.com. 

http://www.msnbc.msn.com/id/19031210