---
module: 085-02
language: en
chapter: 85
title: "Developmental Biology, Germ Layers, Folding, and Placentation"
module_title: "Gastrulation, germ layers, body axes, neurulation, folding, and segmentation"
source_sha256: 1ed6e9281a72d468fff7a576dabf411ed0cf4a6f8d51c67fe9584a4a4b3f5993
---
# Gastrulation to segmentation

## Gastrulation
### Bilaminar disc becomes trilaminar
### Primitive streak appears caudally
### Epiblast cells become motile mesenchyme
#### First ingressing cells form endoderm
#### Next cells form intraembryonic mesoderm
#### Surface cells become ectoderm
### All embryo proper derives from epiblast
### Primitive node is a major organiser
### Notochordal process becomes notochord
#### Defines axis, induces neural development
#### Persists as nucleus pulposus
### Oropharyngeal and cloacal membranes lack mesoderm
### Failed limits: teratoma, caudal regression
#### Midline defects: craniofacial, forebrain

## Germ-layer derivatives
### Surface ectoderm: epidermis, lens, enamel
### Neuroectoderm: neural tube and crest
### Placodes: sensory structures, cranial ganglia
### Mesoderm: connective tissue, muscle, blood, kidney
#### Axial, paraxial, intermediate, lateral plate
### Endoderm: gut, airway, bladder epithelia
#### Liver, pancreas, thyroid, parathyroid, thymus
### Mature organs combine lineages

## Neurulation
### Organiser signals block surface-ectoderm fate
### Neural plate, folds, groove
### Fusion starts near cervical region
### Cranial neuropore first, caudal shortly after
#### Failure: anencephaly to spina bifida
#### Folate lowers but does not abolish risk
### Neural tube becomes CNS and retina
### Alar plate sensory, basal plate motor
#### Sonic hedgehog gradient from floor plate

## Neural crest
### Arises at neural-plate border
### Detaches during closure, migrates widely
### Peripheral neurons, Schwann cells, melanocytes
### Adrenal medulla, craniofacial cartilage and bone
### Conotruncal septation, enteric nervous system
### Defects combine multisystem anomalies

## Mesodermal segmentation
### Somites form in rhythmic sequence
#### Somite number estimates age
### Sclerotome forms vertebrae and ribs
### Myotome forms skeletal muscle
### Dermatome forms dermis of back
### Resegmentation builds vertebral bodies
#### Spinal nerves exit between vertebrae
#### Myotomes bridge adjacent levels
### Intermediate mesoderm: pronephros to metanephros
### Lateral plate splits around coelom
#### Somatopleure: body wall and limbs
#### Splanchnopleure: gut wall and heart
#### Blood islands form endothelium, blood cells

## Axis patterning
### Left-right set near primitive node
#### Nodal cilia drive directional flow
#### NODAL, LEFTY, PITX two on left
#### Disruption: mirror image or heterotaxy
#### Ciliary defects add respiratory disease
### HOX order matches expression domains
### Signals act by concentration and time
#### Same signal, different outcome by history

## Embryonic folding
### Driven by neural tube and somite growth
### Cranial folding moves heart ventrally
#### Foregut incorporated
### Caudal folding incorporates hindgut
### Lateral folds close body wall
#### Except at umbilical ring
### Vitelline duct links midgut to yolk sac
### Defects: omphalocele, gastroschisis, remnants

## Gut, pharynx and timing
### Foregut: coeliac artery
### Midgut: superior mesenteric artery
### Hindgut: inferior mesenteric artery
### Each arch: cartilage, muscle, artery, nerve
#### Pouches form thymus, parathyroids
#### Clefts vanish except external meatus
#### Explain neck cysts and fistulas
### Carnegie stages use morphology
### Embryonic period: high malformation risk
### Trace defects to failed transitions
