Gastrointestinal Tract - Stomach Development

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Developing stomach mid embryonic period (Carnegie stage 13)


This section of notes gives an overview of how the stomach and duodenum develops. The GIT is best imagined as a simple tube, the upper part being the foregut diverticulum, which is further divided into oesophagus and stomach.

During week 4 at the level where the stomach will form the tube begins to dilate, forming an enlarged lumen. The dorsal border grows more rapidly than ventral, which establishes the greater curvature of the stomach.[1] A second rotation (of 90 degrees) occurs on the longitudinal axis establishing the adult orientation of the stomach.

GIT Links: Introduction | Medicine Lecture | Science Lecture | endoderm | mouth | oesophagus | stomach | liver | gallbladder | Pancreas | intestine | mesentery | tongue | taste | enteric nervous system | Stage 13 | Stage 22 | gastrointestinal abnormalities | Movies | Postnatal | milk | tooth | salivary gland | BGD Lecture | BGD Practical | GIT Terms | Category:Gastrointestinal Tract
GIT Histology Links: Upper GIT | Salivary Gland | Smooth Muscle Histology | Liver | Gallbladder | Pancreas | Colon | Histology Stains | Histology | GIT Development
Historic Embryology - Gastrointestinal Tract  
1878 Alimentary Canal | 1882 The Organs of the Inner Germ-Layer The Alimentary Tube with its Appended Organs | 1884 Great omentum and transverse mesocolon | 1902 Meckel's diverticulum | 1902 The Organs of Digestion | 1903 Submaxillary Gland | 1906 Liver | 1907 Development of the Digestive System | 1907 Atlas | 1907 23 Somite Embryo | 1908 Liver | 1908 Liver and Vascular | 1910 Mucous membrane Oesophagus to Small Intestine | 1910 Large intestine and Vermiform process | 1911-13 Intestine and Peritoneum - Part 1 | Part 2 | Part 3 | Part 5 | Part 6 | 1912 Digestive Tract | 1912 Stomach | 1914 Digestive Tract | 1914 Intestines | 1914 Rectum | 1915 Pharynx | 1915 Intestinal Rotation | 1917 Entodermal Canal | 1918 Anatomy | 1921 Alimentary Tube | 1932 Gall Bladder | 1939 Alimentary Canal Looping | 1940 Duodenum anomalies | 2008 Liver | 2016 GIT Notes | Historic Disclaimer
Human Embryo: 1908 13-14 Somite Embryo | 1921 Liver Suspensory Ligament | 1926 22 Somite Embryo | 1907 23 Somite Embryo | 1937 25 Somite Embryo | 1914 27 Somite Embryo | 1914 Week 7 Embryo
Animal Development: 1913 Chicken | 1951 Frog

Historic Embryology: 1912 Stomach

Some Recent Findings

  • Stomach curvature is generated by left-right asymmetric gut morphogenesis[1] "Left-right (LR) asymmetry is a fundamental feature of internal anatomy, yet the emergence of morphological asymmetry remains one of the least understood phases of organogenesis. Asymmetric rotation of the intestine is directed by forces outside the gut, but the morphogenetic events that generate anatomical asymmetry in other regions of the digestive tract remain unknown. Here, we show in mouse and Xenopus that the mechanisms that drive the curvature of the stomach are intrinsic to the gut tube itself. The left wall of the primitive stomach expands more than the right wall, as the left epithelium becomes more polarized and undergoes radial rearrangement. These asymmetries exist across several species, and are dependent on LR patterning genes, including Foxj1, Nodal and Pitx2 Our findings have implications for how LR patterning manifests distinct types of morphological asymmetries in different contexts."
  • FGF4 and retinoic acid direct differentiation of hESCs into PDX1-expressing foregut endoderm in a time- and concentration-dependent manner.[2] "Retinoic acid (RA) and fibroblast growth factor 4 (FGF4) signaling control endoderm patterning and pancreas induction/expansion.FGF
  • FGF10 is required for cell proliferation and gland formation in the stomach epithelium of the chicken embryo. [3] "The development of digestive organs in vertebrates involves active epithelial-mesenchymal interactions. In the chicken proventriculus (glandular stomach), the morphogenesis and cytodifferentiation of the epithelium are controlled by the inductive signaling factors that are secreted from the underlying mesenchyme. ... FGF10 signaling, mediated by FGFR1b and/or FGFR2b, is required for proliferation and gland formation in the epithelium in the developing chick embryo." FGF
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Search term: Stomach Embryology | Stomach Development

Components of Stomach Formation

primitive endoderm

  • foregut diverticulum (pocket)
    • pharyngeal region of foregut
      • laryngo-tracheal groove (see respiratory tract)
      • oesophageal region of foregut
        • oesophagus
          • stomach
            • glandular/proventricular/pyloric stenosis
              • fundus/pyloric antrum
                • pyloric sphincter
            • dorsal mesogastrium
              • lieno-renal ligament
                • splenic primordium
                  • spleen
                • gastro-splenic ligament
                • duodenum (rostral half)
  • foregut-midgut junction
  • midgut region
  • hindgut diverticulum (pocket)

Modified from [4]


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 ‎‎Amniotic Cavity
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 ‎‎Tract Growth
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 ‎‎Stomach Rotation
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 ‎‎Lesser sac
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 ‎‎Greater Omentum
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Stage 13

The images below link to larger cross-sections of the mid-embryonic period (end week 4) stage 13 embryo starting just above the level of the stomach and then in sequence through the stomach to the level of the duodenum.

Stage 13 image 072.jpg Stage 13 image 073.jpg Stage 13 image 074.jpg
D2 Cardio-oesophageal junction D3 Stomach body D4 Stomach body
Stage 13 image 075.jpg Stage 13 image 076.jpg Stage 13 image 077.jpg
D5 Stomach body D6 Pyloric junction D7 Duodenum

 ‎‎GIT Stage 13
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This is an animation based on a reconstruction of the above embryo entire stage 13 gastrointestinal tract.

Links: Carnegie stage 13 - serial sections | Embryo Serial Sections | Movies

Stage 22

The images below link to larger cross-sections of the end of the embryonic period (week 8) stage 22 embryo starting just above the level of the stomach and then in sequence through the stomach to the level of the duodenum. Note how the stomach is "embedded" within the large developing liver.

Stage 22 image 082.jpg Stage 22 image 083.jpg Stage 22 image 084.jpg Stage 22 image 085.jpg
E5 Oesophagus E6 Cardio-oesophageal junction E7 Stomach body Pyloric junction F1 Stomach body Pylorus
Stage 22 image 086.jpg Stage 22 image 087.jpg Stage 22 image 088.jpg Stage 22 image 089.jpg
F2 Stomach body F3 Stomach body Duodenum F4 Duodenal-Jejunal junction F5 Duodenum Jejunum

 ‎‎GIT Stage 22
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This is an animation based on a reconstruction of the above embryo entire stage 22 gastrointestinal tract.

Links: Carnegie stage 22 - serial sections | Embryo Serial Sections | Movies

Stomach Mesentery

In the second trimester, the ventral and dorsal mesenteries associated with the stomach are still anatomically different from the newborn. The figure shows a lateral view of this process comparing the early second trimester arrangement with the newborn structure.

Ventral Mesogastrium

Attached to the superior end of the stomach will form the lesser omentum. This structure will connect the lesser curvature of the stomach to the liver as a ligamentous structure.

Dorsal Mesogastrium

Attached to the inferior end of the stomach initially as an extended fold, this later fuses as a single "apron-like" structure, the greater omentum. Fusion will also incorporate the transverse colon part of the large intestine. This will also contribute the gastrosplenic ligament (gastrolienal ligament).

Greater-omentum.jpg The greater omentum hangs like an apron over the small intestine and transverse colon. It begins attacted to the inferior end of the stomach as a fold of the dorsal mesogastrium which later fuses to form the structure we recognise anatomically. The figure below shows a lateral view of this process comparing the early second trimester arrangement with the newborn structure.

Duodenum/Pancreas Rotation

Pancreas rotation cartoon After the stomach the initial portion of the gastrointestinal tract tube is the duodenum which initially lies in the midline within the peritoneal cavity.

This region, along with the attached pancreas, undergoes rotation to become a retroperitoneal structure.

This diagram shows the rotation with spinal cord at the top, vertebral body then dorsal aorta then pertioneal wall and cavity.


The data below comes from a historic study by Johnson (1910)[5]

  1. Early - A few vacuoles, similar to those of the oesophagus, are found in corresponding stages in the stomach.
  2. 16 mm - the smooth epithelium begins to show a number of pit-like depressions, the first appearances of the gastric pits. These rapidly increase in number and many become elongated to form grooves. At first the basal surface of the epithelium shows no irregularities due to the gastric pits. Then appear slight swellings into the mesenchyma. Still later depressions or furrows, alternating with the gastric pits, extend inward into the epithelium from the mesenchymal surface. This brings about a readjustment of the cells, and causes the heretofore 2 to 3 layered epithelium to become single layered.
  3. The groove-like pits anastomose with one another to form a network. This network marks out irregular areas of the surface epithelium, which have been described as villi.
  4. Growth of the mucous membrane is accompanied by an increase in the number of pits and by an increase in their size. The additional pits develop in between those already formed.
  5. 120 mm - Glands first seen bud out from the bottoms of the pits. These rapidly increase in number and give off branches. Parietal cells are first distinct at 120 mm
  6. Large longitudinal folds of the mucous membrane occur in the stomach, but are variable in position, number, and size. Their occurrence is probably due to the contraction of the muscularis.

Stomach Hormonal Development

The gastrointestinal tract has its own complex entero-endocrine system (enterohormones) that regulates many regional tract functions.

Cells within the stomach express a range of peptide hormones known to regulate a range of gastric functions including secretion of digestive enzymes, mucous and the movement of the luminal contents. The list below shows the earliest detectible presence of specific hormone-containing cells in regions of the developing human stomach.

Hormonal Timecourse

8 weeks - Gastrin containing cells in stomach antrum. Somatostatin cells in both the antrum and the fundus.

10 weeks - Glucagon containing cells in stomach fundus.

11 weeks - Serotonin containing cells in both the antrum and the fundus.

Expression data based upon[6]

Other Gut Peptides

  • Cholecystokinin (CCK), pancreatic polypeptide, peptide YY, glucagon-like peptide-1 (GLP-1), oxyntomodulin (increase satiety and decrease food intake) and ghrelin


The images below show differential gene expression of some selected markers during development (E10.5 and E13.5) of the mouse gastrointestinal tract.[7]



Links: Mouse Development | Full original figure | E10.5 | E13.5


  1. 1.0 1.1 Davis A, Amin NM, Johnson C, Bagley K, Ghashghaei HT & Nascone-Yoder N. (2017). Stomach curvature is generated by left-right asymmetric gut morphogenesis. Development , 144, 1477-1483. PMID: 28242610 DOI.
  2. Johannesson M, Ståhlberg A, Ameri J, Sand FW, Norrman K & Semb H. (2009). FGF4 and retinoic acid direct differentiation of hESCs into PDX1-expressing foregut endoderm in a time- and concentration-dependent manner. PLoS ONE , 4, e4794. PMID: 19277121 DOI.
  3. Shin M, Noji S, Neubüser A & Yasugi S. (2006). FGF10 is required for cell proliferation and gland formation in the stomach epithelium of the chicken embryo. Dev. Biol. , 294, 11-23. PMID: 16616737 DOI.
  4. Kaufman & Bard, The Anatomical Basis of Mouse Development, 1999 Academic Press
  5. Johnson FP. The development of the mucous membrane of the oesophagus, stomach and small intestine in the human embryo. (1910) Amer. J Anat., 10: 521-559.
  6. Stein BA, Buchan AM, Morris J & Polak JM. (1983). The ontogeny of regulatory peptide-containing cells in the human fetal stomach: an immunocytochemical study. J. Histochem. Cytochem. , 31, 1117-25. PMID: 6136542 DOI.
  7. Matsuyama M, Aizawa S & Shimono A. (2009). Sfrp controls apicobasal polarity and oriented cell division in developing gut epithelium. PLoS Genet. , 5, e1000427. PMID: 19300477 DOI.


McCracken KW & Wells JM. (2017). Mechanisms of embryonic stomach development. Semin. Cell Dev. Biol. , 66, 36-42. PMID: 28238948 DOI.

Kim TH & Shivdasani RA. (2016). Stomach development, stem cells and disease. Development , 143, 554-65. PMID: 26884394 DOI.

Zorn AM & Wells JM. (2009). Vertebrate endoderm development and organ formation. Annu. Rev. Cell Dev. Biol. , 25, 221-51. PMID: 19575677 DOI.

Fukuda K & Yasugi S. (2005). The molecular mechanisms of stomach development in vertebrates. Dev. Growth Differ. , 47, 375-82. PMID: 16109035 DOI.

Yasugi S. (1994). Regulation of pepsinogen gene expression in epithelial cells of vertebrate stomach during development. Int. J. Dev. Biol. , 38, 273-9. PMID: 7526882

Johnson LR. (1985). Functional development of the stomach. Annu. Rev. Physiol. , 47, 199-215. PMID: 3922287 DOI.

Deren JS. (1971). Development of structure and function in the fetal and newborn stomach. Am. J. Clin. Nutr. , 24, 144-59. PMID: 4923462


Shin M, Noji S, Neubüser A & Yasugi S. (2006). FGF10 is required for cell proliferation and gland formation in the stomach epithelium of the chicken embryo. Dev. Biol. , 294, 11-23. PMID: 16616737 DOI.

Yasugi S. (2000). Epithelial cell differentiation during stomach development. Hum. Cell , 13, 177-84. PMID: 11329933

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Historic References


Bardeen CR. The critical period in the development of the intestines. (1914) Amer. J Anat. 16: 427 – 445.


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