Gastrointestinal Tract - Intestine Development

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Introduction

midgut herniation

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, jejunum and 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 | gall bladder | Pancreas | intestine | 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 | Gall Bladder | 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 | 1902 The Organs of Digestion | 1903 Submaxillary Gland | 1906 Liver | 1907 Development of the Digestive System | 1907 Atlas | 1907 23 Somite Embryo | 1908 Liver and Vascular | 1910 Mucous membrane Oesophagus to Small Intestine | 1910 Large intestine and Vermiform process | 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 | 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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  • Therefore the list of references do not reflect any editorial selection of material based on content or relevance.
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References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.

Links: References | Discussion Page | Pubmed Most Recent | Journal Searches


Search term: Intestine Embryology

Fatma Uysal, Saffet Ozturk Embryonic poly(A)-binding protein is differently expressed and interacts with the messenger RNAs in the mouse oocytes and early embryos. J. Cell. Biochem.: 2018; PubMed 30302808

Anke Vater, Johann Maierl Adaptive Anatomical Specialization of the Intestines of Alpacas Taking into Account their Original Habitat and Feeding Behaviour. Anat Rec (Hoboken): 2018; PubMed 30288956

Yoonsun Yoon, Kyungju Kim, Suk Keu Yeom, JeeHyun Lee, Yoon Lee A case report of intrahepatic bile duct confluence anomalies in VACTERL syndrome. Medicine (Baltimore): 2018, 97(39);e12411 PubMed 30278516

Mohamed E Abd El-Hack, Dalia H Samak, Ahmed E Noreldin, Karima El-Naggar, Mohamed Abdo Probiotics and plant-derived compounds as eco-friendly agents to inhibit microbial toxins in poultry feed: a comprehensive review. Environ Sci Pollut Res Int: 2018; PubMed 30229484

N Lisnychuk, Yu Soroka, I Andrijchuk, Z Nebesna, K Volkov HISTOLOGICAL CHANGES IN SPLEEN UNDER CONDITIONS OF TOXIC CARCINOGENESIS. Georgian Med News: 2018, (280-281);160-164 PubMed 30204117

Older papers  
  • 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

Adult Small Intestine
Duodenum cartoon.jpg Jejunum and ileum cartoon.jpg
Duodenum Jejunum and ileum
  • Duodenum (adult 25 cm length)
  • Jejunum (adult 1.4 m length)
  • Ileum (adult 3.5 m length)

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

Intestine histology 001.jpg Peyer's patch 01.jpg
Adult jejunum histology Adult ileum Peyer's patches

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

Large Intestine

Large intestine or bowel

  • Cecum (caecum)
    • Vermiform appendix ("appendix", adult 2 to 20 cm length)
  • Colon
    • Ascending colon (adult 25 cm length)
    • Transverse colon
    • Descending colon
    • Sigmoid colon

Intestinal Functions

Small Intestine

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

Cecum

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

Colon

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

Embryonic Development

Week 4

Stage13-GIT-icon.jpg

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)
Stage13-GIT-icon.jpg
 ‎‎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).[7]

Week 8

Stage22-GIT-icon.jpg
Stage22-GIT-icon.jpg
 ‎‎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.[7]

Links: Carnegie stage 22 | Week 8

Rotation

A recent 3 dimensional study[7] 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.[7]

"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[8]

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[9][10]

Fetal rectum growth graph.jpg

Data fromCite error: Invalid <ref> tag; name cannot be a simple integer. Use a descriptive title

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.[11]

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 based upon 8 published reports of necropsy measurement of 1010 guts.[11]

Appendix

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.[12]

There appears to be a developmental anatomical differences in the fetal position of the appendix in males and females.[13] In the fetus, lymphocyte aggregates first appear in this region during the second trimester, week 15 (GA week 17).[14]

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

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

Hindgut

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[15] in human embryos of the Carnegie Collection has been previously carried out (1974). A more recent study[16] 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.[17]


Other studies - [18][19][20][21][22] (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.[23]

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

Abnormalities

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[24] and a later modified by Wallbridge in 1963[25], subsequently two more types of appendix abnormalities have been identified.[26][27]

Modified Cave-Wallbridge Classification (table from[28])

Classification of types
of appendix duplication
Features
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.[29]

  • 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)[30]
  • FGF9

References

  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. 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. 7.0 7.1 7.2 7.3 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.
  8. Martin V & Shaw-Smith C. (2010). Review of genetic factors in intestinal malrotation. Pediatr. Surg. Int. , 26, 769-81. PMID: 20549505 DOI.
  9. FitzSimmons J, Chinn A & Shepard TH. (1988). Normal length of the human fetal gastrointestinal tract. Pediatr Pathol , 8, 633-41. PMID: 3244599
  10. Archie JG, Collins JS & Lebel RR. (2006). Quantitative standards for fetal and neonatal autopsy. Am. J. Clin. Pathol. , 126, 256-65. PMID: 16891202 DOI.
  11. 11.0 11.1 Weaver LT, Austin S & Cole TJ. (1991). Small intestinal length: a factor essential for gut adaptation. Gut , 32, 1321-3. PMID: 1752463
  12. Fisher RE. (2000). The primate appendix: a reassessment. Anat. Rec. , 261, 228-36. PMID: 11135184
  13. 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.
  14. 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.
  15. 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
  16. Hashimoto R. (2013). Development of the human tail bud and splanchnic mesenchyme. Congenit Anom (Kyoto) , 53, 27-33. PMID: 23480355 DOI.
  17. 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.
  18. van der Putte SC. (2009). The development of the human anorectum. Anat Rec (Hoboken) , 292, 951-4. PMID: 19496155 DOI.
  19. Kromer P. (1999). Further study of the urorectal septum in staged human embryos. Folia Morphol. (Warsz) , 58, 53-63. PMID: 10504783
  20. Nievelstein RA, van der Werff JF, Verbeek FJ, Valk J & Vermeij-Keers C. (1998). Normal and abnormal embryonic development of the anorectum in human embryos. Teratology , 57, 70-8. PMID: 9562679 <70::AID-TERA5>3.0.CO;2-A DOI.
  21. Kromer P. (1996). Development of the urorectal septum and differentiation of the urogenital sinus in human embryos of stages 13 to 19. Folia Morphol. (Warsz) , 55, 362-3. PMID: 9243909
  22. Kluth D, Fiegel HC & Metzger R. (2011). Embryology of the hindgut. Semin. Pediatr. Surg. , 20, 152-60. PMID: 21708335 DOI.
  23. 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.
  24. Cave AJ. (1936). Appendix Vermiformis Duplex. J. Anat. , 70, 283-92. PMID: 17104589
  25. WALLBRIDGE PH. (1962). Double appendix. Br J Surg , 50, 346-7. PMID: 13998581
  26. Mesko TW, Lugo R & Breitholtz T. (1989). Horseshoe anomaly of the appendix: a previously undescribed entity. Surgery , 106, 563-6. PMID: 2772830
  27. Tinckler LF. (1968). Triple appendix vermiformis--a unique case. Br J Surg , 55, 79-81. PMID: 5635427
  28. Canbay E & Akman E. (2011). Appendix perforation in appendix duplication in a man: a case report. J Med Case Rep , 5, 162. PMID: 21513538 DOI.
  29. Davì G, Pinto A, Palumbo MG, Gallo V, Mazza A & Strano A. (1985). Dipyridamole and aspirin in arteriosclerosis obliterans of the lower limbs. Adv. Prostaglandin Thromboxane Leukot. Res. , 13, 271-5. PMID: 3159212
  30. Beck F & Stringer EJ. (2010). The role of Cdx genes in the gut and in axial development. Biochem. Soc. Trans. , 38, 353-7. PMID: 20298182 DOI.

Reviews

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

Articles

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

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Cite this page: Hill, M.A. (2018, October 21) Embryology Gastrointestinal Tract - Intestine Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Gastrointestinal_Tract_-_Intestine_Development

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