Gastrointestinal Tract - Intestine Development

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midgut herniation
Intestinal villi and crypts
Intestinal villi and crypts

The part of the gastrointestinal tract (GIT) lying between the stomach and anus, is described as the intestine or bowel. This region is further divided anatomically and functionally into the small intestine or bowel (duodenum, Template:Jejunum and Template:Ileum) and large intestine or bowel (cecum and colon). Initially development concerns the midgut region, connected to the yolk sac, and the hindgut region, ending at the cloacal membrane. This is followed by two mechanical processes of elongation and rotation. Elongation, growth in length, leaves the midgut "herniated" at the umbilicus and external to the abdomen. Rotation, around a mesentery axis, establishes the anatomical position of the large intestine within the peritoneal space.

Migration of neural crest cells into the wall establishes the enteric nervous system, which has a role in peristalsis and secretion. Prenatally, secretions also accumulate in this region and are the first postnatal bowel movement, the meconium.

The small intestine grows in length rapidly in the last trimester, at birth it is about half the eventual adult length (More? Small Intestine Length). Like most of the gut, this region is not "functional" until after birth, when development continues by populating the large intestine with commensal bacteria and the establishment of the immune structure in the wall.

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 Small Intestine | 1912 large Intestine | 1915 Intestinal Rotation

Some Recent Findings

Model for cloacal septation[1]
  • Radial WNT5A-Guided Post-mitotic Filopodial Pathfinding Is Critical for Midgut Tube Elongation[2] "The early midgut undergoes intensive elongation, but the underlying cellular and molecular mechanisms are unknown. The early midgut epithelium is pseudostratified, and its nuclei travel between apical and basal surfaces in concert with cell cycle. Using 3D confocal imaging and 2D live imaging, we profiled behaviors of individual dividing cells. As nuclei migrate apically for mitosis, cells maintain a basal process (BP), which splits but is inherited by only one daughter. After mitosis, some daughters directly use the inherited BP as a "conduit" to transport the nucleus basally, while >50% of daughters generate a new basal filopodium and use it as a path to return the nucleus. Post-mitotic filopodial "pathfinding" is guided by mesenchymal WNT5A. Without WNT5A, some cells fail to tether basally and undergo apoptosis, leading to a shortened midgut. Thus, these studies reveal previously unrecognized strategies for efficient post-mitotic nuclear trafficking, which is critical for early midgut elongation." WNT
  • Growth of the colon and rectum throughout gestation: evaluation with fetal MRI[3] "Congenital abnormalities of the gastrointestinal tract are increasingly being evaluated by prenatal magnetic resonance imaging (MRI). However, there is a paucity of reports describing the normal quantitative development of the fetal colon and rectum on MRI. This study provides normal ranges of the prenatal colon and rectum as a function of gestational age. They may serve as reference values when interpreting fetal MRI."
  • Digestive Tract in Human Embryos Between Carnegie Stages 11 and 13[4] "We selected 37 human embryos at Carnegie Stage (CS) 11-13 (28-33 days after fertilization) and three-dimensionally analyzed the morphology and positioning of the digestive tract and derived primordia in all samples, using images reconstructed from histological serial sections. The digestive tract was initially formed by a narrowing of the yolk sac, and then several derived primordia such as the pharynx, lung, stomach, liver, and dorsal pancreas primordia differentiated during CS12 (21-29 somites) and CS13 (≥ 30 somites). The differentiation of four pairs of pharyngeal pouches was complete in all CS13 embryos. The respiratory primordium was recognized in ≥ 26-somite embryos and it flattened and then branched at CS13. The trachea formed and then elongated in ≥ 35-somite embryos. The stomach adopted a spindle shape in all ≥ 34-somite embryos, and the liver bud was recognized in ≥ 27-somite embryos. The dorsal pancreas appeared as definitive buddings in all but three CS13 embryos, and around these buddings, the small intestine bent in ≥ 33-somite embryos. In ≥ 35-somite embryos, the small intestine rotated around the cranial-caudal axis and had begun to form a primitive intestinal loop, which led to umbilical herniation."
More recent papers  
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Search term: Intestine Embryology | Intestine Development | Small Intestine Development | Large Intestine Development |Duodenum Development | Jejunum Development | Ileum Development | Appendix Development | Colon Development

Older papers  
These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table.

See also the Discussion Page for other references listed by year and References on this current page.

  • Review - How to make an intestine[5] With the high prevalence of gastrointestinal disorders, there is great interest in establishing in vitro models of human intestinal disease and in developing drug-screening platforms that more accurately represent the complex physiology of the intestine. We will review how recent advances in developmental and stem cell biology have made it possible to generate complex, three-dimensional, human intestinal tissues in vitro through directed differentiation of human pluripotent stem cells.
  • Bmp7 functions via a polarity mechanism to promote cloacal septation[1] "During normal development in human and other placental mammals, the embryonic cloacal cavity separates along the axial longitudinal plane to give rise to the urethral system, ventrally, and the rectum, dorsally. Defects in cloacal development are very common and present clinically as a rectourethral fistula in about 1 in 5,000 live human births. Yet, the cellular mechanisms of cloacal septation remain poorly understood. ...Our results strongly indicate that Bmp7/JNK signaling regulates remodeling of the cloacal endoderm resulting in a topological separation of the urinary and digestive systems. Our study points to the importance of Bmp and JNK signaling in cloacal development and rectourethral malformations."
  • Fgf9 signaling regulates small intestinal elongation and mesenchymal development[6] "Short bowel syndrome is an acquired condition in which the length of the small intestine is insufficient to perform its normal absorptive function. ...These data suggest a model in which epithelial-derived Fgf9 stimulates intestinal mesenchymal stem cells (iMSCs) that in turn regulate underlying mesenchymal fibroblast proliferation and differentiation at least in part through inhibition of Tgfbeta signaling in the mesenchyme."

Small Intestine

The small intestine (or bowel) consists of 3 main anatomical regions. A notable feature of this epithelium is the continuous replacement throughout life, from a stem cell population residing in the crypts, migrating towards the tips of the villi.

Adult Length

  • Duodenum (25 cm)
  • Jejunum (1.4 m)
  • Ileum (3.5 m)



The duodenum develops from the part of the tube immediately after the stomach and uniquely forms the junction between foregut (upper) and midgut (lower). Unlike the remainder of the small intestine it grows significantly less in overall length. Initially suspended by a mesentery, and projects forward in the form of a loop, both of which rotate with the fusion of the two pancreatic buds, and subsequently displaced by the transverse colon. The right surface of the duodenal mesentery is directed backward to parietal peritoneum and becomes retroperitoneal. Both the biliary and the pancreatic ducts empty into the duodenum.

Late Embryonic Duodenum (Week 8, GA week 10, Stage 22)
Stage 22 image 186.jpg Note the distinctive "star-shaped" lumen of duodenum and the developing muscular wall.

The pancreas and its duct can also be seen on the right of image.

Search PubMed: Duodenum Development


Jejunum and ileum
Jejunum and ileum

The part of the small intestine between the duodenum and ileum. A recent study has shown that the postnatal jejunal lumen contains a distinctive bacterial population (facultative anaerobes and oxygen-tolerant obligate anaerobes) similar to those found in the oral cavity though with additional enterobacteriaceae (Gram-negative bacteria).[7]

Adult jejunum histology

Intestine histology 001.jpg

Search PubMed: Jejunum Development


The adult ileum contains specialised aggregated lymphoid nodules known as Peyer's patches.

Adult ileum Peyer's patches

Peyer's patch 01.jpg

See small intestine or bowel length (see also Fetal Intestine Length and Small Intestine Length)

Search PubMed: Ileum Development

Large Intestine

Development of the large intestine and its mesentery
Lymphatics of Cecum and Vermiform Process

Large intestine or bowel is where water is absorbed from the indigestible food residue and where the majority of the enteric microbiota reside. Before birth the intestine is sterile, after birth the microbiota is established from the mother. Later postnatally by diet, lifestyle and antibiotic exposure. The large intestine bacteria is mainly anaerobic, compared to the aerobic found in other regions.


(Latin, caecum) forms the beginning of the large intestine.

Vermiform Appendix

("appendix", adult 2 to 20 cm length) The vermiform appendix is a highly immune structure and can act as a "microbiota reservoir" during antibiotic and other treatment.


Anatomical Parts

  1. Ascending colon (adult 25 cm length)
  2. Transverse colon
  3. Descending colon
  4. Sigmoid colon

Search PubMed: Large Intestine Development | Cecum Intestine Development | Vermiform appendix Intestine Development | Colon Development |

Intestinal Functions

Small Intestine

  • absorption of nutrients and minerals found in food
  • Duodenum - principal site for iron absorption


  • connects the ileum with the ascending colon
  • separated by the ileocecal valve (ICV, Bauhin's valve)
  • connected to the vermiform appendix ("appendix")


  • absorbs fluid, water and salts, from solid wastes
  • site of commensal bacteria (flora) fermentation of unabsorbed material

Embryonic Development

Week 4


Colour code:

  • Foregut - oropharyngeal membrane, oesophagus, pharynx (dark salmon - Foregut, oropharyngeal membrane, oesophagus, pharynx)
  • Trachea and Lung Buds (dark blue - Trachea and Lung Buds)
  • Mesonephros (yellow - mesonephros)
  • Mesonephric Ducts (light blue - mesonephric ducts)
  • Hindgut and Cloaca (brown - hindgut and cloaca)
 ‎‎GIT Stage 13
Page | Play

Week 7

Small intestine secondary loops week 7 to 8

Human embryo small intestine secondary loops (week 7 to 8).[8]

Week 8

 ‎‎GIT Stage 22
Page | Play

Stage 22 image 088.jpg Stage 22 image 089.jpg

Late embryonic small intestine commencing at the duodenum, continuing as ventrally herniated and returning to join the colon.

Small intestine tertiary loops week 8

Small intestine tertiary loops week 8.[8]

Links: Carnegie stage 22 | Week 8


A recent 3 dimensional study[8] has suggested a modified “en-bloc rotation” of the small intestine, compared the the earlier simplified description of 270 degree rotation (below).

Human intestine “en-bloc rotation” model.jpg

Human intestine “en-bloc rotation” model.[8]

"If one insists on using the term rotation for this movement, it would be largely around a craniocaudal axis (in the transverse plane) rather than a dorsoventral axis (frontal plane). In view of the brief time window and orientation of the apparent rotational axis, we conclude that the distal ileum and cecum “slide” rather than “rotate” as from the umbilical orifice to the lower-right abdominal cavity."

Normal intestinal rotation cartoon.jpg

Normal intestinal rotation[9]

Fetal Intestine Length

Fetal small Intestine length growth graph.jpg Fetal large Intestine length growth graph.jpg
Fetal small Intestine length growth Fetal Large Intestine length growth

Data from[10][11]

Fetal rectum growth graph.jpg

Data from[3]

Small Intestine Length

Small intestine growth in length is initially linear (first half pregnancy to 32 cm CRL), followed by rapid growth in the last 15 weeks doubling the overall length. Growth continues postnatally but after 1 year slows again to a linear increase to adulthood.[12]

Small Intestine Length
Age (weeks gestational age) Average Length (cm)
20 125
30 200
term 275
1 year postnatal 380
5 years 450
10 years 500
20 years 575
Table data from 8 published reports of necropsy measurement of 1010 guts.[12]
Links: Gastrointestinal Tract - Postnatal


Appendix (2nd month of fetal life)[13]

The appendix (vermiform appendix, vermix) is a finger-like diverticulum located anatomically at the cecum, the junction between the small and large intestines (colon). The length (2.5-13 cm) is longer in both infants and children and also has more abundant lymphatic tissue in early life. The wall structure is similar to the small intestine (though with no villi), nor plicae circularis. Its immune function is associated with the many lymph nodules surrounding the lumen that extend from the mucosa into the submucosa. It has also been suggested as a repository for beneficial intestinal microflora. See also the review comparing the appendix in different species.[14]

There appears to be a developmental anatomical differences in the fetal position of the appendix in males and females.[15] In the fetus, lymphocyte aggregates first appear in this region during the second trimester, week 15 (GA week 17).[16]

Historically, Berengario da Carpi (1460-c.1530) in Commentaria cum amplissimis additionibus super Anatomia Mundini (1521) was the first to describe the human appendix.

See also notes on appendix duplication.

Links: Immune System Development | Lecture - Lymphatic Structure and Organs | Search PubMed


Anatomically the distal third of the transverse colon and the splenic flexure, the descending colon, sigmoid colon and rectum. The developmental timing of the anus and rectum formation[17] in human embryos of the Carnegie Collection has been previously carried out (1974). A more recent study[18] has also been made of the Kyoto Collection embryos.

There has been some recent controversy over the "anal membrane" formation.

A recent study hindgut and anorectum development in human embryos shows that WNT5a is active in this region prior to anus formation, when it is down-regulated.[19]

Other studies - [20][21][22][23][24] (rat)

Intestinal Motility

The enteric nervous system neural crest-derived neurons interacts with the circular and longitudinal smooth muscle layers and the interstitial cells of Cajal to generate motility. The developmental timing data shown below is from a recent review.[25]

Neural Crest

week 5 - migrating neural crest cells reach the midgut

week 7 - neural crest cells have colonized the entire gut

  • colonization occurs in a rostro-caudal sequence

Gastrointestinal Tract Plexuses (enteric nervous system)
Myenteric plexus Submucosal plexus
Auerbach's plexus Meissner's plexus
Leopold Auerbach (1828–1897) a German anatomist and neuropathologist. Georg Meissner (1829–1905) a German anatomist and physiologist.
  • first formed plexus
  • lies between the outer longitudinal and inner circular smooth muscle layers of muscularis externa
  • provides motor innervation to both layers
  • secretomotor innervation to the mucosa
  • both parasympathetic and sympathetic input
  • forms 2-3 days after the myenteric plexus
  • formed by cells migrating from the myenteric plexus
  • innervates smooth muscle of the muscularis mucosae
  • only parasympathetic fibers
  Links: enteric nervous system | intestine | neural crest | PMID 25428846

Smooth Muscle

week 8 - esophagus circular muscle

week 11 - hindgut circular muscle

week 14 - hindgut concentric muscularis mucosae, circular muscle, and longitudinal muscle

Interstitial Cells of Cajal

Interstitial cells of Cajal (ICC) are electrical pacemaker cells within the gastrointestinal tract smooth muscle. They create the basal (slow waves) rhythm required for contraction and peristalsis. They are mesodermal in origin.

weeks 7-9 - cells initially appear

week 11 - distinct clusters

week 12-14 - clustered around myenteric ganglia along the entire gut

Links: Neural Crest Development


Abnormality Links: Gastrointestinal Tract - Abnormalities | Intestine Development | Gastrointestinal Tract
Links: Gastrointestinal Tract - Abnormalities | Image - Small intestine duplication

Appendix Duplication

Appendix duplication is an extremely rare congenital anomaly (0.004% to 0.009% of appendectomy specimens) first classified according to their anatomic location by Cave in 1936[26] and a later modified by Wallbridge in 1963[27], subsequently two more types of appendix abnormalities have been identified.[28][29]

Modified Cave-Wallbridge Classification (table from[30])

Classification of types
of appendix duplication
A Single cecum with various degrees of incomplete duplication
B1 (bird type) Two appendixes symmetrically placed on either side of the ileocecal valve
B2 (tenia coli type) ne appendix arises from the cecum at the usual site, and the second

appendix branches from the cecum along the lines of the tenia at various distances from the first

B3 One appendix arises from the usual site, and the second appendix arises from

the hepatic flexura

B4 One appendix arises from the usual site, and the second appendix arises from

the splenic flexura

C Double cecum, each with an appendix
Horseshoe appendix One appendix has two openings into a common cecum
Triple appendix One appendix arises from the cecum at the usual site, and two additional appendixes arise from the colon

Short Bowel Syndrome

Short bowel syndrome (SBS) results typically due to developmental abnormalities, extensive intestinal resection during the neonatal period, or necrotising enterolitis.[31]

  • reduces gut function for digestion and absorption of nutrients (intestinal failure).

Links: PubMed Health | Better Health

Molecular Factors

  • Cdx (Caudal-type homeobox) group of ParaHox genes (mouse Cdx1, Cdx2 and Cdx4)[32]
  • FGF9


  1. 1.0 1.1 Xu K, Wu X, Shapiro E, Huang H, Zhang L, Hickling D, Deng Y, Lee P, Li J, Lepor H & Grishina I. (2012). Bmp7 functions via a polarity mechanism to promote cloacal septation. PLoS ONE , 7, e29372. PMID: 22253716 DOI.
  2. Wang S, Cebrian C, Schnell S & Gumucio DL. (2018). Radial WNT5A-Guided Post-mitotic Filopodial Pathfinding Is Critical for Midgut Tube Elongation. Dev. Cell , 46, 173-188.e3. PMID: 30016620 DOI.
  3. 3.0 3.1 Ben-Nun MS, Ben-Shlush A & Raviv Zilka L. (2018). Growth of the colon and rectum throughout gestation: evaluation with fetal MRI. Acta Radiol Open , 7, 2058460118761206. PMID: 29531795 DOI.
  4. Ueno S, Yamada S, Uwabe C, Männer J, Shiraki N & Takakuwa T. (2016). The Digestive Tract and Derived Primordia Differentiate by Following a Precise Timeline in Human Embryos Between Carnegie Stages 11 and 13. Anat Rec (Hoboken) , 299, 439-49. PMID: 26995337 DOI.
  5. Wells JM & Spence JR. (2014). How to make an intestine. Development , 141, 752-60. PMID: 24496613 DOI.
  6. Geske MJ, Zhang X, Patel KK, Ornitz DM & Stappenbeck TS. (2008). Fgf9 signaling regulates small intestinal elongation and mesenchymal development. Development , 135, 2959-68. PMID: 18653563 DOI.
  7. Sundin OH, Mendoza-Ladd A, Zeng M, Diaz-Arévalo D, Morales E, Fagan BM, Ordoñez J, Velez P, Antony N & McCallum RW. (2017). The human jejunum has an endogenous microbiota that differs from those in the oral cavity and colon. BMC Microbiol. , 17, 160. PMID: 28716079 DOI.
  8. 8.0 8.1 8.2 8.3 Soffers JH, Hikspoors JP, Mekonen HK, Koehler SE & Lamers WH. (2015). The growth pattern of the human intestine and its mesentery. BMC Dev. Biol. , 15, 31. PMID: 26297675 DOI.
  9. Martin V & Shaw-Smith C. (2010). Review of genetic factors in intestinal malrotation. Pediatr. Surg. Int. , 26, 769-81. PMID: 20549505 DOI.
  10. FitzSimmons J, Chinn A & Shepard TH. (1988). Normal length of the human fetal gastrointestinal tract. Pediatr Pathol , 8, 633-41. PMID: 3244599
  11. Archie JG, Collins JS & Lebel RR. (2006). Quantitative standards for fetal and neonatal autopsy. Am. J. Clin. Pathol. , 126, 256-65. PMID: 16891202 DOI.
  12. 12.0 12.1 Weaver LT, Austin S & Cole TJ. (1991). Small intestinal length: a factor essential for gut adaptation. Gut , 32, 1321-3. PMID: 1752463
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  14. Fisher RE. (2000). The primate appendix: a reassessment. Anat. Rec. , 261, 228-36. PMID: 11135184
  15. Malas MA, Gökçimen A & Sulak O. (2001). Growing of caecum and vermiform appendix during the fetal period. Fetal. Diagn. Ther. , 16, 173-7. PMID: 11316934 DOI.
  16. Malas MA, Sulak O, Gökçimen A & Sari A. (2004). Development of the vermiform appendix during the fetal period. Surg Radiol Anat , 26, 202-7. PMID: 15173960 DOI.
  17. de Vries PA. and Friedland GW. The staged sequential development of the anus and rectum in human embryos and fetuses. (1974) J. Pediatr. Surg., 9(5): 755-69 PMID 4424274
  18. Hashimoto R. (2013). Development of the human tail bud and splanchnic mesenchyme. Congenit Anom (Kyoto) , 53, 27-33. PMID: 23480355 DOI.
  19. Li FF, Zhang T, Bai YZ, Yuan ZW & Wang WL. (2011). Spatiotemporal expression of Wnt5a during the development of the hindgut and anorectum in human embryos. Int J Colorectal Dis , 26, 983-8. PMID: 21431850 DOI.
  20. van der Putte SC. (2009). The development of the human anorectum. Anat Rec (Hoboken) , 292, 951-4. PMID: 19496155 DOI.
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Kostouros A, Koliarakis I, Natsis K, Spandidos DA, Tsatsakis A & Tsiaoussis J. (2020). Large intestine embryogenesis: Molecular pathways and related disorders (Review). Int. J. Mol. Med. , , . PMID: 32319546 DOI.

Chin AM, Hill DR, Aurora M & Spence JR. (2017). Morphogenesis and maturation of the embryonic and postnatal intestine. Semin. Cell Dev. Biol. , 66, 81-93. PMID: 28161556 DOI.

Viotti M, Foley AC & Hadjantonakis AK. (2014). Gutsy moves in mice: cellular and molecular dynamics of endoderm morphogenesis. Philos. Trans. R. Soc. Lond., B, Biol. Sci. , 369, . PMID: 25349455 DOI.

Wells JM & Spence JR. (2014). How to make an intestine. Development , 141, 752-60. PMID: 24496613 DOI.

Shyer AE, Tallinen T, Nerurkar NL, Wei Z, Gil ES, Kaplan DL, Tabin CJ & Mahadevan L. (2013). Villification: how the gut gets its villi. Science , 342, 212-8. PMID: 23989955 DOI.

Noah TK, Donahue B & Shroyer NF. (2011). Intestinal development and differentiation. Exp. Cell Res. , 317, 2702-10. PMID: 21978911 DOI.

Burns AJ, Roberts RR, Bornstein JC & Young HM. (2009). Development of the enteric nervous system and its role in intestinal motility during fetal and early postnatal stages. Semin. Pediatr. Surg. , 18, 196-205. PMID: 19782301 DOI.


Cho BH, Kim JH, Jin ZW, Wilting J, Rodríguez-Vázquez JF & Murakami G. (2018). Topographical anatomy of the intestines during in utero physiological herniation. Clin Anat , 31, 583-592. PMID: 29044646 DOI.

Soffers JH, Hikspoors JP, Mekonen HK, Koehler SE & Lamers WH. (2015). The growth pattern of the human intestine and its mesentery. BMC Dev. Biol. , 15, 31. PMID: 26297675 DOI.

Ueda Y, Yamada S, Uwabe C, Kose K & Takakuwa T. (2016). Intestinal Rotation and Physiological Umbilical Herniation During the Embryonic Period. Anat Rec (Hoboken) , 299, 197-206. PMID: 26599074 DOI.

Kim TH, Kim BM, Mao J, Rowan S & Shivdasani RA. (2011). Endodermal Hedgehog signals modulate Notch pathway activity in the developing digestive tract mesenchyme. Development , 138, 3225-33. PMID: 21750033 DOI.

Kim WK, Kim H, Ahn DH, Kim MH & Park HW. (2003). Timetable for intestinal rotation in staged human embryos and fetuses. Birth Defects Res. Part A Clin. Mol. Teratol. , 67, 941-5. PMID: 14745932 DOI.

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Historic Disclaimer - information about historic embryology pages 
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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

Tandler J. The development of the human duodenum in early embryonic stages. (Zur Entwicklungsgeschichte des menschlichen Duodenum in frṻhen Embryonalstadien) (1900) Morphol. Jahrb. 29:187-216.

Tandler (1900) was the first to recognize that the lumen of the duodenum in embryos between " 30 to 60 days," is normally "more or less completely" obliterated.

Frazer JE. and Robbins RH. On the factors concerned in causing rotation of the intestine in man. (1915) J Anat. 50(1): 75-110. PMID 17233053

Cave AJ. Appendix vermiformis duplex. (1936) J Anat. 70: 283-292. PMID 171045891

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