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Gastrointestinal tract

The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and other animals, including the esophagus, stomach, and intestines. Food taken in through the mouth is digested to extract nutrients and absorb energy, and the waste expelled at the anus as faeces. Gastrointestinal is an adjective meaning of or pertaining to the stomach and intestines.

Gastrointestinal tract
Diagram of stomach, intestines and rectum in the average human
Details
SystemDigestive system
Identifiers
Latintractus digestorius (mouth to anus),
canalis alimentarius (esophagus to large intestine),
canalis gastrointestinales stomach to large intestine)
MeSHD041981
Anatomical terminology
[edit on Wikidata]

Most animals have a "through-gut" or complete digestive tract. Exceptions are more primitive ones: sponges have small pores (ostia) throughout their body for digestion and a larger dorsal pore (osculum) for excretion, comb jellies have both a ventral mouth and dorsal anal pores, while cnidarians and acoels have a single pore for both digestion and excretion.[1][2]

The human gastrointestinal tract consists of the esophagus, stomach, and intestines, and is divided into the upper and lower gastrointestinal tracts.[3] The GI tract includes all structures between the mouth and the anus,[4] forming a continuous passageway that includes the main organs of digestion, namely, the stomach, small intestine, and large intestine. The complete human digestive system is made up of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver and gallbladder).[5] The tract may also be divided into foregut, midgut, and hindgut, reflecting the embryological origin of each segment. The whole human GI tract is about nine meters (30 feet) long at autopsy. It is considerably shorter in the living body because the intestines, which are tubes of smooth muscle tissue, maintain constant muscle tone in a halfway-tense state but can relax in spots to allow for local distention and peristalsis.[6][7]

The gastrointestinal tract contains the gut microbiota, with some 1,000 different strains of bacteria having diverse roles in the maintenance of immune health and metabolism, and many other microorganisms.[8][9][10] Cells of the GI tract release hormones to help regulate the digestive process. These digestive hormones, including gastrin, secretin, cholecystokinin, and ghrelin, are mediated through either intracrine or autocrine mechanisms, indicating that the cells releasing these hormones are conserved structures throughout evolution.[11]

Human gastrointestinal tract edit

Structure edit

 Salivary glandsParotid glandSubmandibular glandSublingual glandpharynxTongueEsophagusPancreasStomachPancreatic ductIleumAnusRectumVermiform appendixCecumDescending colonAscending colonTransverse colonColon (anatomy)Bile ductDuodenumGallbladderLiveroral cavity
Upper and lower human gastrointestinal tract

The structure and function can be described both as gross anatomy and as microscopic anatomy or histology. The tract itself is divided into upper and lower tracts, and the intestines small and large parts.[12]

Upper gastrointestinal tract edit

The upper gastrointestinal tract consists of the mouth, pharynx, esophagus, stomach, and duodenum.[13] The exact demarcation between the upper and lower tracts is the suspensory muscle of the duodenum. This differentiates the embryonic borders between the foregut and midgut, and is also the division commonly used by clinicians to describe gastrointestinal bleeding as being of either "upper" or "lower" origin. Upon dissection, the duodenum may appear to be a unified organ, but it is divided into four segments based on function, location, and internal anatomy. The four segments of the duodenum are as follows (starting at the stomach, and moving toward the jejunum): bulb, descending, horizontal, and ascending. The suspensory muscle attaches the superior border of the ascending duodenum to the diaphragm.

The suspensory muscle is an important anatomical landmark that shows the formal division between the duodenum and the jejunum, the first and second parts of the small intestine, respectively.[14] This is a thin muscle which is derived from the embryonic mesoderm.

Lower gastrointestinal tract edit

The lower gastrointestinal tract includes most of the small intestine and all of the large intestine.[15] In human anatomy, the intestine (bowel, or gut. Greek: éntera) is the segment of the gastrointestinal tract extending from the pyloric sphincter of the stomach to the anus and as in other mammals, consists of two segments: the small intestine and the large intestine. In humans, the small intestine is further subdivided into the duodenum, jejunum, and ileum while the large intestine is subdivided into the cecum, ascending, transverse, descending, and sigmoid colon, rectum, and anal canal.[16][17]

Small intestine edit
 
Illustration of the small intestine

The small intestine begins at the duodenum and is a tubular structure, usually between 6 and 7 m long.[18] Its mucosal area in an adult human is about 30 m2 (320 sq ft).[19] The combination of the circular folds, the villi, and the microvilli increases the absorptive area of the mucosa about 600-fold, making a total area of about 250 m2 (2,700 sq ft) for the entire small intestine.[20] Its main function is to absorb the products of digestion (including carbohydrates, proteins, lipids, and vitamins) into the bloodstream. There are three major divisions:

  1. Duodenum: A short structure (about 20–25 cm long[18]) that receives chyme from the stomach, together with pancreatic juice containing digestive enzymes and bile from the gall bladder. The digestive enzymes break down proteins, and bile emulsifies fats into micelles. The duodenum contains Brunner's glands which produce a mucus-rich alkaline secretion containing bicarbonate. These secretions, in combination with bicarbonate from the pancreas, neutralize the stomach acids contained in the chyme.
  2. Jejunum: This is the midsection of the small intestine, connecting the duodenum to the ileum. It is about 2.5 m (8.2 ft) long and contains the circular folds also known as plicae circulares and villi that increase its surface area. Products of digestion (sugars, amino acids, and fatty acids) are absorbed into the bloodstream here.
  3. Ileum: The final section of the small intestine. It is about 3 m long, and contains villi similar to the jejunum. It absorbs mainly vitamin B12 and bile acids, as well as any other remaining nutrients.
Large intestine edit

The large intestine, also called the colon, forms an arch starting at the cecum and ending at the rectum and anal canal. It also includes the appendix, which is attached to the cecum. Its length is about 1.5 m, and the area of the mucosa in an adult human is about 2 m2 (22 sq ft).[19] Its main function is to absorb water and salts. The colon is further divided into:

  1. Cecum (first portion of the colon) and appendix
  2. Ascending colon (ascending in the back wall of the abdomen)
  3. Right colic flexure (flexed portion of the ascending and transverse colon apparent to the liver)
  4. Transverse colon (passing below the diaphragm)
  5. Left colic flexure (flexed portion of the transverse and descending colon apparent to the spleen)
  6. Descending colon (descending down the left side of the abdomen)
  7. Sigmoid colon (a loop of the colon closest to the rectum)
  8. Rectum
  9. Anal canal

Development edit

The gut is an endoderm-derived structure. At approximately the sixteenth day of human development, the embryo begins to fold ventrally (with the embryo's ventral surface becoming concave) in two directions: the sides of the embryo fold in on each other and the head and tail fold toward one another. The result is that a piece of the yolk sac, an endoderm-lined structure in contact with the ventral aspect of the embryo, begins to be pinched off to become the primitive gut. The yolk sac remains connected to the gut tube via the vitelline duct. Usually, this structure regresses during development; in cases where it does not, it is known as Meckel's diverticulum.

During fetal life, the primitive gut is gradually patterned into three segments: foregut, midgut, and hindgut. Although these terms are often used in reference to segments of the primitive gut, they are also used regularly to describe regions of the definitive gut as well.

Each segment of the gut is further specified and gives rise to specific gut and gut-related structures in later development. Components derived from the gut proper, including the stomach and colon, develop as swellings or dilatations in the cells of the primitive gut. In contrast, gut-related derivatives — that is, those structures that derive from the primitive gut but are not part of the gut proper, in general, develop as out-pouchings of the primitive gut. The blood vessels supplying these structures remain constant throughout development.[21]

Part Part in adult Gives rise to Arterial supply
Foregut esophagus to first 2 sections of the duodenum Esophagus, stomach, duodenum (1st and 2nd parts), liver, gallbladder, pancreas, superior portion of pancreas
(Though the spleen is supplied by the celiac trunk, it is derived from dorsal mesentery and therefore not a foregut derivative)
celiac trunk
Midgut lower duodenum, to the first two-thirds of the transverse colon lower duodenum, jejunum, ileum, cecum, appendix, ascending colon, and first two-thirds of the transverse colon branches of the superior mesenteric artery
Hindgut last third of the transverse colon, to the upper part of the anal canal last third of the transverse colon, descending colon, rectum, and upper part of the anal canal branches of the inferior mesenteric artery

Histology edit

 
General structure of the gut wall
  • 1: Mucosa: Epithelium
  • 2: Mucosa: Lamina propria
  • 3: Mucosa: Muscularis mucosae
  • 4: Lumen
  • 5: Lymphatic tissue
  • 6: Duct of gland outside tract
  • 7: Gland in mucosa
  • 8: Submucosa
  • 9: Glands in submucosa
  • 10: Meissner's submucosal plexus
  • 11: Vein
  • 12: Muscularis: Circular muscle
  • 13: Muscularis: Longitudinal muscle
  • 14: Serosa: Areolar connective tissue
  • 15: Serosa: Epithelium
  • 16: Auerbach's myenteric plexus
  • 17: Nerve
  • 18: Artery
  • 19: Mesentery

The gastrointestinal tract has a form of general histology with some differences that reflect the specialization in functional anatomy.[22] The GI tract can be divided into four concentric layers in the following order:

Mucosa edit

The mucosa is the innermost layer of the gastrointestinal tract. The mucosa surrounds the lumen, or open space within the tube. This layer comes in direct contact with digested food (chyme). The mucosa is made up of:

  • Epithelium – innermost layer. Responsible for most digestive, absorptive and secretory processes.
  • Lamina propria – a layer of connective tissue. Unusually cellular compared to most connective tissue
  • Muscularis mucosae – a thin layer of smooth muscle that aids the passing of material and enhances the interaction between the epithelial layer and the contents of the lumen by agitation and peristalsis

The mucosae are highly specialized in each organ of the gastrointestinal tract to deal with the different conditions. The most variation is seen in the epithelium.

Submucosa edit

The submucosa consists of a dense irregular layer of connective tissue with large blood vessels, lymphatics, and nerves branching into the mucosa and muscularis externa. It contains the submucosal plexus, an enteric nervous plexus, situated on the inner surface of the muscularis externa.

Muscular layer edit

The muscular layer consists of an inner circular layer and a longitudinal outer layer. The circular layer prevents food from traveling backward and the longitudinal layer shortens the tract. The layers are not truly longitudinal or circular, rather the layers of muscle are helical with different pitches. The inner circular is helical with a steep pitch and the outer longitudinal is helical with a much shallower pitch.[23] Whilst the muscularis externa is similar throughout the entire gastrointestinal tract, an exception is the stomach which has an additional inner oblique muscular layer to aid with grinding and mixing of food. The muscularis externa of the stomach is composed of the inner oblique layer, middle circular layer, and outer longitudinal layer.

Between the circular and longitudinal muscle layers is the myenteric plexus. This controls peristalsis. Activity is initiated by the pacemaker cells, (myenteric interstitial cells of Cajal). The gut has intrinsic peristaltic activity (basal electrical rhythm) due to its self-contained enteric nervous system. The rate can be modulated by the rest of the autonomic nervous system.[23]

The coordinated contractions of these layers is called peristalsis and propels the food through the tract. Food in the GI tract is called a bolus (ball of food) from the mouth down to the stomach. After the stomach, the food is partially digested and semi-liquid, and is referred to as chyme. In the large intestine, the remaining semi-solid substance is referred to as faeces.[23]

Adventitia and serosa edit

The outermost layer of the gastrointestinal tract consists of several layers of connective tissue.

Intraperitoneal parts of the GI tract are covered with serosa. These include most of the stomach, first part of the duodenum, all of the small intestine, caecum and appendix, transverse colon, sigmoid colon and rectum. In these sections of the gut, there is a clear boundary between the gut and the surrounding tissue. These parts of the tract have a mesentery.

Retroperitoneal parts are covered with adventitia. They blend into the surrounding tissue and are fixed in position. For example, the retroperitoneal section of the duodenum usually passes through the transpyloric plane. These include the esophagus, pylorus of the stomach, distal duodenum, ascending colon, descending colon and anal canal. In addition, the oral cavity has adventitia.

Gene and protein expression edit

Approximately 20,000 protein coding genes are expressed in human cells and 75% of these genes are expressed in at least one of the different parts of the digestive organ system.[24][25] Over 600 of these genes are more specifically expressed in one or more parts of the GI tract and the corresponding proteins have functions related to digestion of food and uptake of nutrients. Examples of specific proteins with such functions are pepsinogen PGC and the lipase LIPF, expressed in chief cells, and gastric ATPase ATP4A and gastric intrinsic factor GIF, expressed in parietal cells of the stomach mucosa. Specific proteins expressed in the stomach and duodenum involved in defence include mucin proteins, such as mucin 6 and intelectin-1.[26]

Transit time edit

The time taken for food to transit through the gastrointestinal tract varies on multiple factors, including age, ethnicity, and gender.[medical citation needed] Several techniques have been used to measure transit time, including radiography following a barium-labeled meal, breath hydrogen analysis, scintigraphic analysis following a radiolabeled meal,[27] and simple ingestion and spotting of corn kernels.[28] It takes 2.5 to 3 hours for 50% of the contents to leave the stomach.[medical citation needed] The rate of digestion is also dependent of the material being digested, as food composition from the same meal may leave the stomach at different rates.[medical citation needed] Total emptying of the stomach takes around 4–5 hours, and transit through the colon takes 30 to 50 hours.[27][29][30]

Immune function edit

The gastrointestinal tract forms an important part of the immune system.[31]

Immune barrier edit

The surface area of the digestive tract is estimated to be about 32 square meters, or about half a badminton court.[19] With such a large exposure (more than three times larger than the exposed surface of the skin), these immune components function to prevent pathogens from entering the blood and lymph circulatory systems.[32] Fundamental components of this protection are provided by the intestinal mucosal barrier, which is composed of physical, biochemical, and immune elements elaborated by the intestinal mucosa.[33] Microorganisms also are kept at bay by an extensive immune system comprising the gut-associated lymphoid tissue (GALT)

There are additional factors contributing to protection from pathogen invasion. For example, low pH (ranging from 1 to 4) of the stomach is fatal for many microorganisms that enter it.[34] Similarly, mucus (containing IgA antibodies) neutralizes many pathogenic microorganisms.[35] Other factors in the GI tract contribution to immune function include enzymes secreted in the saliva and bile.

Immune system homeostasis edit

Beneficial bacteria also can contribute to the homeostasis of the gastrointestinal immune system. For example, Clostridia, one of the most predominant bacterial groups in the GI tract, play an important role in influencing the dynamics of the gut's immune system.[36] It has been demonstrated that the intake of a high fiber diet could be responsible for the induction of T-regulatory cells (Tregs). This is due to the production of short-chain fatty acids during the fermentation of plant-derived nutrients such as butyrate and propionate. Basically, the butyrate induces the differentiation of Treg cells by enhancing histone H3 acetylation in the promoter and conserved non-coding sequence regions of the FOXP3 locus, thus regulating the T cells, resulting in the reduction of the inflammatory response and allergies.

Intestinal microbiota edit

The large intestine contains multiple types of bacteria that can break down molecules the human body cannot process alone,[37] demonstrating a symbiotic relationship. These bacteria are responsible for gas production at host–pathogen interface, which is released as flatulence. However, the primary function of the large intestine is water absorption from digested material (regulated by the hypothalamus) and the reabsorption of sodium and nutrients.[citation needed]

Beneficial intestinal bacteria compete with potentially harmful bacteria for space and "food", as the intestinal tract has limited resources. A ratio of 80–85% beneficial to 15–20% potentially harmful bacteria is proposed for maintaining homeostasis.[citation needed] An imbalanced ratio results in dysbiosis.

Detoxification and drug metabolism edit

Enzymes such as CYP3A4, along with the antiporter activities, are also instrumental in the intestine's role of drug metabolism in the detoxification of antigens and xenobiotics.[38]

Other animals edit

In most vertebrates, including fishes, amphibians, birds, reptiles, and egg-laying mammals, the gastrointestinal tract ends in a cloaca and not an anus. In the cloaca, the urinary system is fused with the genito-anal pore. Therians (all mammals that do not lay eggs, including humans) possess separate anal and uro-genital openings. The females of the subgroup placentalia have even separate urinary and genital openings.

During early development the asymmetric position of the bowels and inner organs is initiated (see also axial twist theory).

Ruminants show many specializations for digesting and fermenting tough plant material, consisting of additional stomach compartments.

Many birds and other animals have a specialised stomach in the digestive tract called a gizzard used for grinding up food.

Another feature found in a range of animals is the crop. In birds this is found as a pouch alongside the esophagus.

In 2020, the oldest known fossil digestive tract, of an extinct wormlike organism in the Cloudinidae was discovered; it lived during the late Ediacaran period about 550 million years ago.[39][40]

A through-gut (one with both mouth and anus) is thought to have evolved within the nephrozoan clade of Bilateria, after their ancestral ventral orifice (single, as in cnidarians and acoels; re-evolved in nephrozoans like flatworms) stretched antero-posteriorly, before the middle part of the stretch would get narrower and closed fully, leaving an anterior orifice (mouth) and a posterior orifice (anus plus genital opening). A stretched gut without the middle part closed is present in another branch of bilaterians, the extinct proarticulates. This and the amphistomic development (when both mouth and anus develop from the gut stretch in the embryo) present in some nephrozoans (e.g. roundworms) are considered to support this hypothesis.[41][42]

Clinical significance edit

Diseases edit

There are many diseases and conditions that can affect the gastrointestinal system, including infections, inflammation and cancer.

Various pathogens, such as bacteria that cause foodborne illnesses, can induce gastroenteritis which results from inflammation of the stomach and small intestine. Antibiotics to treat such bacterial infections can decrease the microbiome diversity of the gastrointestinal tract, and further enable inflammatory mediators.[43] Gastroenteritis is the most common disease of the GI tract.

Diverticular disease is a condition that is very common in older people in industrialized countries. It usually affects the large intestine but has been known to affect the small intestine as well. Diverticulosis occurs when pouches form on the intestinal wall. Once the pouches become inflamed it is known as diverticulitis.

Inflammatory bowel disease is an inflammatory condition affecting the bowel walls, and includes the subtypes Crohn's disease and ulcerative colitis. While Crohn's can affect the entire gastrointestinal tract, ulcerative colitis is limited to the large intestine. Crohn's disease is widely regarded as an autoimmune disease. Although ulcerative colitis is often treated as though it were an autoimmune disease, there is no consensus that it actually is such.

Functional gastrointestinal disorders the most common of which is irritable bowel syndrome. Functional constipation and chronic functional abdominal pain are other functional disorders of the intestine that have physiological causes but do not have identifiable structural, chemical, or infectious pathologies.

Symptoms edit

Several symptoms can indicate problems with the gastrointestinal tract, including:

Treatment edit

Gastrointestinal surgery can often be performed in the outpatient setting. In the United States in 2012, operations on the digestive system accounted for 3 of the 25 most common ambulatory surgery procedures and constituted 9.1 percent of all outpatient ambulatory surgeries.[44]

Imaging edit

Various methods of imaging the gastrointestinal tract include the upper and lower gastrointestinal series:

Other related diseases edit

  • Cholera
  • Enteric duplication cyst
  • Giardiasis
  • Pancreatitis
  • Peptic ulcer disease
  • Yellow fever
  • Helicobacter pylori is a gram-negative spiral bacterium. Over half the world's population is infected with it, mainly during childhood; it is not certain how the disease is transmitted. It colonizes the gastrointestinal system, predominantly the stomach. The bacterium has specific survival conditions that are specific to the human gastric microenvironment: it is both capnophilic and microaerophilic. Helicobacter also exhibits a tropism for gastric epithelial lining and the gastric mucosal layer about it. Gastric colonization of this bacterium triggers a robust immune response leading to moderate to severe inflammation, known as gastritis. Signs and symptoms of infection are gastritis, burning abdominal pain, weight loss, loss of appetite, bloating, burping, nausea, bloody vomit, and black tarry stools. Infection can be detected in a number of ways: GI X-rays, endoscopy, blood tests for anti-Helicobacter antibodies, a stool test, and a urease breath test (which is a by-product of the bacteria). If caught soon enough, it can be treated with three doses of different proton pump inhibitors as well as two antibiotics, taking about a week to cure. If not caught soon enough, surgery may be required.[45][46][47][48]
  • Intestinal pseudo-obstruction is a syndrome caused by a malformation of the digestive system, characterized by a severe impairment in the ability of the intestines to push and assimilate. Symptoms include daily abdominal and stomach pain, nausea, severe distension, vomiting, heartburn, dysphagia, diarrhea, constipation, dehydration and malnutrition. There is no cure for intestinal pseudo-obstruction. Different types of surgery and treatment managing life-threatening complications such as ileus and volvulus, intestinal stasis which lead to bacterial overgrowth, and resection of affected or dead parts of the gut may be needed. Many patients require parenteral nutrition.
  • Ileus is a blockage of the intestines.
  • Coeliac disease is a common form of malabsorption, affecting up to 1% of people of northern European descent. An autoimmune response is triggered in intestinal cells by digestion of gluten proteins. Ingestion of proteins found in wheat, barley and rye, causes villous atrophy in the small intestine. Lifelong dietary avoidance of these foodstuffs in a gluten-free diet is the only treatment.
  • Enteroviruses are named by their transmission-route through the intestine (enteric meaning intestinal), but their symptoms are not mainly associated with the intestine.
  • Endometriosis can affect the intestines, with similar symptoms to IBS.
  • Bowel twist (or similarly, bowel strangulation) is a comparatively rare event (usually developing sometime after major bowel surgery). It is, however, hard to diagnose correctly, and if left uncorrected can lead to bowel infarction and death. (The singer Maurice Gibb is understood to have died from this.)
  • Angiodysplasia of the colon
  • Constipation
  • Diarrhea
  • Hirschsprung's disease (aganglionosis)
  • Intussusception
  • Polyp (medicine) (see also colorectal polyp)
  • Pseudomembranous colitis
  • Toxic megacolon usually a complication of ulcerative colitis

Uses of animal guts edit

Intestines from animals other than humans are used in a number of ways. From each species of livestock that is a source of milk, a corresponding rennet is obtained from the intestines of milk-fed calves. Pig and calf intestines are eaten, and pig intestines are used as sausage casings. Calf intestines supply calf-intestinal alkaline phosphatase (CIP), and are used to make goldbeater's skin. Other uses are:

  • The use of animal gut strings by musicians can be traced back to the third dynasty of Egypt. In the recent past, strings were made out of lamb gut. With the advent of the modern era, musicians have tended to use strings made of silk, or synthetic materials such as nylon or steel. Some instrumentalists, however, still use gut strings in order to evoke the older tone quality. Although such strings were commonly referred to as "catgut" strings, cats were never used as a source for gut strings.[49]
  • Sheep gut was the original source for natural gut string used in racquets, such as for tennis. Today, synthetic strings are much more common, but the best gut strings are now made out of cow gut.
  • Gut cord has also been used to produce strings for the snares that provide a snare drum's characteristic buzzing timbre. While the modern snare drum almost always uses metal wire rather than gut cord, the North African bendir frame drum still uses gut for this purpose.
  • "Natural" sausage hulls, or casings, are made of animal gut, especially hog, beef, and lamb.
  • The wrapping of kokoretsi, gardoubakia, and torcinello is made of lamb (or goat) gut.
  • Haggis is traditionally boiled in, and served in, a sheep stomach.
  • Chitterlings, a kind of food, consist of thoroughly washed pig's gut.
  • Animal gut was used to make the cord lines in longcase clocks and for fusee movements in bracket clocks, but may be replaced by metal wire.
  • The oldest known condoms, from 1640 AD, were made from animal intestine.[50]

See also edit

References edit

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External links edit

  • The gastro intestinal tract in the Human Protein Atlas

gastrointestinal, tract, enteric, redirects, here, other, uses, enteric, disambiguation, gastrointestinal, tract, tract, digestive, tract, alimentary, canal, tract, passageway, digestive, system, that, leads, from, mouth, anus, tract, contains, major, organs, . Enteric redirects here For other uses see Enteric disambiguation The gastrointestinal tract GI tract digestive tract alimentary canal is the tract or passageway of the digestive system that leads from the mouth to the anus The GI tract contains all the major organs of the digestive system in humans and other animals including the esophagus stomach and intestines Food taken in through the mouth is digested to extract nutrients and absorb energy and the waste expelled at the anus as faeces Gastrointestinal is an adjective meaning of or pertaining to the stomach and intestines Gastrointestinal tractDiagram of stomach intestines and rectum in the average humanDetailsSystemDigestive systemIdentifiersLatintractus digestorius mouth to anus canalis alimentarius esophagus to large intestine canalis gastrointestinales stomach to large intestine MeSHD041981Anatomical terminology edit on Wikidata Most animals have a through gut or complete digestive tract Exceptions are more primitive ones sponges have small pores ostia throughout their body for digestion and a larger dorsal pore osculum for excretion comb jellies have both a ventral mouth and dorsal anal pores while cnidarians and acoels have a single pore for both digestion and excretion 1 2 The human gastrointestinal tract consists of the esophagus stomach and intestines and is divided into the upper and lower gastrointestinal tracts 3 The GI tract includes all structures between the mouth and the anus 4 forming a continuous passageway that includes the main organs of digestion namely the stomach small intestine and large intestine The complete human digestive system is made up of the gastrointestinal tract plus the accessory organs of digestion the tongue salivary glands pancreas liver and gallbladder 5 The tract may also be divided into foregut midgut and hindgut reflecting the embryological origin of each segment The whole human GI tract is about nine meters 30 feet long at autopsy It is considerably shorter in the living body because the intestines which are tubes of smooth muscle tissue maintain constant muscle tone in a halfway tense state but can relax in spots to allow for local distention and peristalsis 6 7 The gastrointestinal tract contains the gut microbiota with some 1 000 different strains of bacteria having diverse roles in the maintenance of immune health and metabolism and many other microorganisms 8 9 10 Cells of the GI tract release hormones to help regulate the digestive process These digestive hormones including gastrin secretin cholecystokinin and ghrelin are mediated through either intracrine or autocrine mechanisms indicating that the cells releasing these hormones are conserved structures throughout evolution 11 Contents 1 Human gastrointestinal tract 1 1 Structure 1 1 1 Upper gastrointestinal tract 1 1 2 Lower gastrointestinal tract 1 1 2 1 Small intestine 1 1 2 2 Large intestine 1 1 3 Development 1 1 4 Histology 1 1 4 1 Mucosa 1 1 4 2 Submucosa 1 1 4 3 Muscular layer 1 1 4 4 Adventitia and serosa 1 1 5 Gene and protein expression 1 1 6 Transit time 1 1 7 Immune function 1 1 7 1 Immune barrier 1 1 7 2 Immune system homeostasis 1 1 8 Intestinal microbiota 1 1 9 Detoxification and drug metabolism 2 Other animals 3 Clinical significance 3 1 Diseases 3 2 Symptoms 3 3 Treatment 3 4 Imaging 3 5 Other related diseases 4 Uses of animal guts 5 See also 6 References 7 External linksHuman gastrointestinal tract editStructure edit nbsp Upper and lower human gastrointestinal tractThe structure and function can be described both as gross anatomy and as microscopic anatomy or histology The tract itself is divided into upper and lower tracts and the intestines small and large parts 12 Upper gastrointestinal tract edit Main articles Mouth Pharynx Esophagus Stomach and duodenum The upper gastrointestinal tract consists of the mouth pharynx esophagus stomach and duodenum 13 The exact demarcation between the upper and lower tracts is the suspensory muscle of the duodenum This differentiates the embryonic borders between the foregut and midgut and is also the division commonly used by clinicians to describe gastrointestinal bleeding as being of either upper or lower origin Upon dissection the duodenum may appear to be a unified organ but it is divided into four segments based on function location and internal anatomy The four segments of the duodenum are as follows starting at the stomach and moving toward the jejunum bulb descending horizontal and ascending The suspensory muscle attaches the superior border of the ascending duodenum to the diaphragm The suspensory muscle is an important anatomical landmark that shows the formal division between the duodenum and the jejunum the first and second parts of the small intestine respectively 14 This is a thin muscle which is derived from the embryonic mesoderm Lower gastrointestinal tract edit Guts redirects here For other uses see Gut disambiguation The lower gastrointestinal tract includes most of the small intestine and all of the large intestine 15 In human anatomy the intestine bowel or gut Greek entera is the segment of the gastrointestinal tract extending from the pyloric sphincter of the stomach to the anus and as in other mammals consists of two segments the small intestine and the large intestine In humans the small intestine is further subdivided into the duodenum jejunum and ileum while the large intestine is subdivided into the cecum ascending transverse descending and sigmoid colon rectum and anal canal 16 17 Small intestine edit nbsp Illustration of the small intestineMain articles Small intestine Duodenum Jejunum and Ileum The small intestine begins at the duodenum and is a tubular structure usually between 6 and 7 m long 18 Its mucosal area in an adult human is about 30 m2 320 sq ft 19 The combination of the circular folds the villi and the microvilli increases the absorptive area of the mucosa about 600 fold making a total area of about 250 m2 2 700 sq ft for the entire small intestine 20 Its main function is to absorb the products of digestion including carbohydrates proteins lipids and vitamins into the bloodstream There are three major divisions Duodenum A short structure about 20 25 cm long 18 that receives chyme from the stomach together with pancreatic juice containing digestive enzymes and bile from the gall bladder The digestive enzymes break down proteins and bile emulsifies fats into micelles The duodenum contains Brunner s glands which produce a mucus rich alkaline secretion containing bicarbonate These secretions in combination with bicarbonate from the pancreas neutralize the stomach acids contained in the chyme Jejunum This is the midsection of the small intestine connecting the duodenum to the ileum It is about 2 5 m 8 2 ft long and contains the circular folds also known as plicae circulares and villi that increase its surface area Products of digestion sugars amino acids and fatty acids are absorbed into the bloodstream here Ileum The final section of the small intestine It is about 3 m long and contains villi similar to the jejunum It absorbs mainly vitamin B12 and bile acids as well as any other remaining nutrients Large intestine edit Main article Large intestine The large intestine also called the colon forms an arch starting at the cecum and ending at the rectum and anal canal It also includes the appendix which is attached to the cecum Its length is about 1 5 m and the area of the mucosa in an adult human is about 2 m2 22 sq ft 19 Its main function is to absorb water and salts The colon is further divided into Cecum first portion of the colon and appendix Ascending colon ascending in the back wall of the abdomen Right colic flexure flexed portion of the ascending and transverse colon apparent to the liver Transverse colon passing below the diaphragm Left colic flexure flexed portion of the transverse and descending colon apparent to the spleen Descending colon descending down the left side of the abdomen Sigmoid colon a loop of the colon closest to the rectum Rectum Anal canalDevelopment edit Main article Development of the digestive system The gut is an endoderm derived structure At approximately the sixteenth day of human development the embryo begins to fold ventrally with the embryo s ventral surface becoming concave in two directions the sides of the embryo fold in on each other and the head and tail fold toward one another The result is that a piece of the yolk sac an endoderm lined structure in contact with the ventral aspect of the embryo begins to be pinched off to become the primitive gut The yolk sac remains connected to the gut tube via the vitelline duct Usually this structure regresses during development in cases where it does not it is known as Meckel s diverticulum During fetal life the primitive gut is gradually patterned into three segments foregut midgut and hindgut Although these terms are often used in reference to segments of the primitive gut they are also used regularly to describe regions of the definitive gut as well Each segment of the gut is further specified and gives rise to specific gut and gut related structures in later development Components derived from the gut proper including the stomach and colon develop as swellings or dilatations in the cells of the primitive gut In contrast gut related derivatives that is those structures that derive from the primitive gut but are not part of the gut proper in general develop as out pouchings of the primitive gut The blood vessels supplying these structures remain constant throughout development 21 Part Part in adult Gives rise to Arterial supplyForegut esophagus to first 2 sections of the duodenum Esophagus stomach duodenum 1st and 2nd parts liver gallbladder pancreas superior portion of pancreas Though the spleen is supplied by the celiac trunk it is derived from dorsal mesentery and therefore not a foregut derivative celiac trunkMidgut lower duodenum to the first two thirds of the transverse colon lower duodenum jejunum ileum cecum appendix ascending colon and first two thirds of the transverse colon branches of the superior mesenteric arteryHindgut last third of the transverse colon to the upper part of the anal canal last third of the transverse colon descending colon rectum and upper part of the anal canal branches of the inferior mesenteric arteryHistology edit Main article Gastrointestinal wall nbsp General structure of the gut wall 1 Mucosa Epithelium2 Mucosa Lamina propria3 Mucosa Muscularis mucosae4 Lumen5 Lymphatic tissue6 Duct of gland outside tract7 Gland in mucosa8 Submucosa9 Glands in submucosa10 Meissner s submucosal plexus11 Vein12 Muscularis Circular muscle13 Muscularis Longitudinal muscle14 Serosa Areolar connective tissue15 Serosa Epithelium16 Auerbach s myenteric plexus17 Nerve18 Artery19 MesenteryThe gastrointestinal tract has a form of general histology with some differences that reflect the specialization in functional anatomy 22 The GI tract can be divided into four concentric layers in the following order Mucosa Submucosa Muscular layer Adventitia or serosaMucosa edit See also Oral mucosa and Gastric mucosa The mucosa is the innermost layer of the gastrointestinal tract The mucosa surrounds the lumen or open space within the tube This layer comes in direct contact with digested food chyme The mucosa is made up of Epithelium innermost layer Responsible for most digestive absorptive and secretory processes Lamina propria a layer of connective tissue Unusually cellular compared to most connective tissue Muscularis mucosae a thin layer of smooth muscle that aids the passing of material and enhances the interaction between the epithelial layer and the contents of the lumen by agitation and peristalsisThe mucosae are highly specialized in each organ of the gastrointestinal tract to deal with the different conditions The most variation is seen in the epithelium Submucosa edit Main article Submucosa The submucosa consists of a dense irregular layer of connective tissue with large blood vessels lymphatics and nerves branching into the mucosa and muscularis externa It contains the submucosal plexus an enteric nervous plexus situated on the inner surface of the muscularis externa Muscular layer edit The muscular layer consists of an inner circular layer and a longitudinal outer layer The circular layer prevents food from traveling backward and the longitudinal layer shortens the tract The layers are not truly longitudinal or circular rather the layers of muscle are helical with different pitches The inner circular is helical with a steep pitch and the outer longitudinal is helical with a much shallower pitch 23 Whilst the muscularis externa is similar throughout the entire gastrointestinal tract an exception is the stomach which has an additional inner oblique muscular layer to aid with grinding and mixing of food The muscularis externa of the stomach is composed of the inner oblique layer middle circular layer and outer longitudinal layer Between the circular and longitudinal muscle layers is the myenteric plexus This controls peristalsis Activity is initiated by the pacemaker cells myenteric interstitial cells of Cajal The gut has intrinsic peristaltic activity basal electrical rhythm due to its self contained enteric nervous system The rate can be modulated by the rest of the autonomic nervous system 23 The coordinated contractions of these layers is called peristalsis and propels the food through the tract Food in the GI tract is called a bolus ball of food from the mouth down to the stomach After the stomach the food is partially digested and semi liquid and is referred to as chyme In the large intestine the remaining semi solid substance is referred to as faeces 23 Adventitia and serosa edit Main articles Serous membrane and Adventitia The outermost layer of the gastrointestinal tract consists of several layers of connective tissue Intraperitoneal parts of the GI tract are covered with serosa These include most of the stomach first part of the duodenum all of the small intestine caecum and appendix transverse colon sigmoid colon and rectum In these sections of the gut there is a clear boundary between the gut and the surrounding tissue These parts of the tract have a mesentery Retroperitoneal parts are covered with adventitia They blend into the surrounding tissue and are fixed in position For example the retroperitoneal section of the duodenum usually passes through the transpyloric plane These include the esophagus pylorus of the stomach distal duodenum ascending colon descending colon and anal canal In addition the oral cavity has adventitia Gene and protein expression edit Approximately 20 000 protein coding genes are expressed in human cells and 75 of these genes are expressed in at least one of the different parts of the digestive organ system 24 25 Over 600 of these genes are more specifically expressed in one or more parts of the GI tract and the corresponding proteins have functions related to digestion of food and uptake of nutrients Examples of specific proteins with such functions are pepsinogen PGC and the lipase LIPF expressed in chief cells and gastric ATPase ATP4A and gastric intrinsic factor GIF expressed in parietal cells of the stomach mucosa Specific proteins expressed in the stomach and duodenum involved in defence include mucin proteins such as mucin 6 and intelectin 1 26 Transit time edit The time taken for food to transit through the gastrointestinal tract varies on multiple factors including age ethnicity and gender medical citation needed Several techniques have been used to measure transit time including radiography following a barium labeled meal breath hydrogen analysis scintigraphic analysis following a radiolabeled meal 27 and simple ingestion and spotting of corn kernels 28 It takes 2 5 to 3 hours for 50 of the contents to leave the stomach medical citation needed The rate of digestion is also dependent of the material being digested as food composition from the same meal may leave the stomach at different rates medical citation needed Total emptying of the stomach takes around 4 5 hours and transit through the colon takes 30 to 50 hours 27 29 30 Immune function edit The gastrointestinal tract forms an important part of the immune system 31 Immune barrier edit The surface area of the digestive tract is estimated to be about 32 square meters or about half a badminton court 19 With such a large exposure more than three times larger than the exposed surface of the skin these immune components function to prevent pathogens from entering the blood and lymph circulatory systems 32 Fundamental components of this protection are provided by the intestinal mucosal barrier which is composed of physical biochemical and immune elements elaborated by the intestinal mucosa 33 Microorganisms also are kept at bay by an extensive immune system comprising the gut associated lymphoid tissue GALT There are additional factors contributing to protection from pathogen invasion For example low pH ranging from 1 to 4 of the stomach is fatal for many microorganisms that enter it 34 Similarly mucus containing IgA antibodies neutralizes many pathogenic microorganisms 35 Other factors in the GI tract contribution to immune function include enzymes secreted in the saliva and bile Immune system homeostasis edit Beneficial bacteria also can contribute to the homeostasis of the gastrointestinal immune system For example Clostridia one of the most predominant bacterial groups in the GI tract play an important role in influencing the dynamics of the gut s immune system 36 It has been demonstrated that the intake of a high fiber diet could be responsible for the induction of T regulatory cells Tregs This is due to the production of short chain fatty acids during the fermentation of plant derived nutrients such as butyrate and propionate Basically the butyrate induces the differentiation of Treg cells by enhancing histone H3 acetylation in the promoter and conserved non coding sequence regions of the FOXP3 locus thus regulating the T cells resulting in the reduction of the inflammatory response and allergies Intestinal microbiota edit The large intestine contains multiple types of bacteria that can break down molecules the human body cannot process alone 37 demonstrating a symbiotic relationship These bacteria are responsible for gas production at host pathogen interface which is released as flatulence However the primary function of the large intestine is water absorption from digested material regulated by the hypothalamus and the reabsorption of sodium and nutrients citation needed Beneficial intestinal bacteria compete with potentially harmful bacteria for space and food as the intestinal tract has limited resources A ratio of 80 85 beneficial to 15 20 potentially harmful bacteria is proposed for maintaining homeostasis citation needed An imbalanced ratio results in dysbiosis Detoxification and drug metabolism edit Enzymes such as CYP3A4 along with the antiporter activities are also instrumental in the intestine s role of drug metabolism in the detoxification of antigens and xenobiotics 38 Other animals editIn most vertebrates including fishes amphibians birds reptiles and egg laying mammals the gastrointestinal tract ends in a cloaca and not an anus In the cloaca the urinary system is fused with the genito anal pore Therians all mammals that do not lay eggs including humans possess separate anal and uro genital openings The females of the subgroup placentalia have even separate urinary and genital openings During early development the asymmetric position of the bowels and inner organs is initiated see also axial twist theory Ruminants show many specializations for digesting and fermenting tough plant material consisting of additional stomach compartments Many birds and other animals have a specialised stomach in the digestive tract called a gizzard used for grinding up food Another feature found in a range of animals is the crop In birds this is found as a pouch alongside the esophagus In 2020 the oldest known fossil digestive tract of an extinct wormlike organism in the Cloudinidae was discovered it lived during the late Ediacaran period about 550 million years ago 39 40 A through gut one with both mouth and anus is thought to have evolved within the nephrozoan clade of Bilateria after their ancestral ventral orifice single as in cnidarians and acoels re evolved in nephrozoans like flatworms stretched antero posteriorly before the middle part of the stretch would get narrower and closed fully leaving an anterior orifice mouth and a posterior orifice anus plus genital opening A stretched gut without the middle part closed is present in another branch of bilaterians the extinct proarticulates This and the amphistomic development when both mouth and anus develop from the gut stretch in the embryo present in some nephrozoans e g roundworms are considered to support this hypothesis 41 42 Clinical significance editThis section discusses related diseases medical associations with the gastrointestinal tract and use in surgery Main articles Gastrointestinal disease and Gastroenterology Further information Clinical significance Diseases edit There are many diseases and conditions that can affect the gastrointestinal system including infections inflammation and cancer Various pathogens such as bacteria that cause foodborne illnesses can induce gastroenteritis which results from inflammation of the stomach and small intestine Antibiotics to treat such bacterial infections can decrease the microbiome diversity of the gastrointestinal tract and further enable inflammatory mediators 43 Gastroenteritis is the most common disease of the GI tract Gastrointestinal cancer may occur at any point in the gastrointestinal tract and includes mouth cancer tongue cancer oesophageal cancer stomach cancer and colorectal cancer Inflammatory conditions Ileitis is an inflammation of the ileum colitis is an inflammation of the large intestine Appendicitis is inflammation of the appendix located at the caecum This is a potentially fatal condition if left untreated most cases of appendicitis require surgical intervention Diverticular disease is a condition that is very common in older people in industrialized countries It usually affects the large intestine but has been known to affect the small intestine as well Diverticulosis occurs when pouches form on the intestinal wall Once the pouches become inflamed it is known as diverticulitis Inflammatory bowel disease is an inflammatory condition affecting the bowel walls and includes the subtypes Crohn s disease and ulcerative colitis While Crohn s can affect the entire gastrointestinal tract ulcerative colitis is limited to the large intestine Crohn s disease is widely regarded as an autoimmune disease Although ulcerative colitis is often treated as though it were an autoimmune disease there is no consensus that it actually is such Functional gastrointestinal disorders the most common of which is irritable bowel syndrome Functional constipation and chronic functional abdominal pain are other functional disorders of the intestine that have physiological causes but do not have identifiable structural chemical or infectious pathologies Symptoms edit Several symptoms can indicate problems with the gastrointestinal tract including Vomiting which may include regurgitation of food or the vomiting of blood Diarrhea or the passage of liquid or more frequent stools Constipation which refers to the passage of fewer and hardened stools Blood in stool which includes fresh red blood maroon coloured blood and tarry coloured bloodTreatment edit Gastrointestinal surgery can often be performed in the outpatient setting In the United States in 2012 operations on the digestive system accounted for 3 of the 25 most common ambulatory surgery procedures and constituted 9 1 percent of all outpatient ambulatory surgeries 44 Imaging edit Various methods of imaging the gastrointestinal tract include the upper and lower gastrointestinal series Radioopaque dyes may be swallowed to produce a barium swallow Parts of the tract may be visualised by camera This is known as endoscopy if examining the upper gastrointestinal tract and colonoscopy or sigmoidoscopy if examining the lower gastrointestinal tract Capsule endoscopy is where a capsule containing a camera is swallowed in order to examine the tract Biopsies may also be taken when examined An abdominal x ray may be used to examine the lower gastrointestinal tract Other related diseases edit Cholera Enteric duplication cyst Giardiasis Pancreatitis Peptic ulcer disease Yellow fever Helicobacter pylori is a gram negative spiral bacterium Over half the world s population is infected with it mainly during childhood it is not certain how the disease is transmitted It colonizes the gastrointestinal system predominantly the stomach The bacterium has specific survival conditions that are specific to the human gastric microenvironment it is both capnophilic and microaerophilic Helicobacter also exhibits a tropism for gastric epithelial lining and the gastric mucosal layer about it Gastric colonization of this bacterium triggers a robust immune response leading to moderate to severe inflammation known as gastritis Signs and symptoms of infection are gastritis burning abdominal pain weight loss loss of appetite bloating burping nausea bloody vomit and black tarry stools Infection can be detected in a number of ways GI X rays endoscopy blood tests for anti Helicobacter antibodies a stool test and a urease breath test which is a by product of the bacteria If caught soon enough it can be treated with three doses of different proton pump inhibitors as well as two antibiotics taking about a week to cure If not caught soon enough surgery may be required 45 46 47 48 Intestinal pseudo obstruction is a syndrome caused by a malformation of the digestive system characterized by a severe impairment in the ability of the intestines to push and assimilate Symptoms include daily abdominal and stomach pain nausea severe distension vomiting heartburn dysphagia diarrhea constipation dehydration and malnutrition There is no cure for intestinal pseudo obstruction Different types of surgery and treatment managing life threatening complications such as ileus and volvulus intestinal stasis which lead to bacterial overgrowth and resection of affected or dead parts of the gut may be needed Many patients require parenteral nutrition Ileus is a blockage of the intestines Coeliac disease is a common form of malabsorption affecting up to 1 of people of northern European descent An autoimmune response is triggered in intestinal cells by digestion of gluten proteins Ingestion of proteins found in wheat barley and rye causes villous atrophy in the small intestine Lifelong dietary avoidance of these foodstuffs in a gluten free diet is the only treatment Enteroviruses are named by their transmission route through the intestine enteric meaning intestinal but their symptoms are not mainly associated with the intestine Endometriosis can affect the intestines with similar symptoms to IBS Bowel twist or similarly bowel strangulation is a comparatively rare event usually developing sometime after major bowel surgery It is however hard to diagnose correctly and if left uncorrected can lead to bowel infarction and death The singer Maurice Gibb is understood to have died from this Angiodysplasia of the colon Constipation Diarrhea Hirschsprung s disease aganglionosis Intussusception Polyp medicine see also colorectal polyp Pseudomembranous colitis Toxic megacolon usually a complication of ulcerative colitisUses of animal guts editIntestines from animals other than humans are used in a number of ways From each species of livestock that is a source of milk a corresponding rennet is obtained from the intestines of milk fed calves Pig and calf intestines are eaten and pig intestines are used as sausage casings Calf intestines supply calf intestinal alkaline phosphatase CIP and are used to make goldbeater s skin Other uses are The use of animal gut strings by musicians can be traced back to the third dynasty of Egypt In the recent past strings were made out of lamb gut With the advent of the modern era musicians have tended to use strings made of silk or synthetic materials such as nylon or steel Some instrumentalists however still use gut strings in order to evoke the older tone quality Although such strings were commonly referred to as catgut strings cats were never used as a source for gut strings 49 Sheep gut was the original source for natural gut string used in racquets such as for tennis Today synthetic strings are much more common but the best gut strings are now made out of cow gut Gut cord has also been used to produce strings for the snares that provide a snare drum s characteristic buzzing timbre While the modern snare drum almost always uses metal wire rather than gut cord the North African bendir frame drum still uses gut for this purpose Natural sausage hulls or casings are made of animal gut especially hog beef and lamb The wrapping of kokoretsi gardoubakia and torcinello is made of lamb or goat gut Haggis is traditionally boiled in and served in a sheep stomach Chitterlings a kind of food consist of thoroughly washed pig s gut Animal gut was used to make the cord lines in longcase clocks and for fusee movements in bracket clocks but may be replaced by metal wire The oldest known condoms from 1640 AD were made from animal intestine 50 See also editThis article uses anatomical terminology Gastrointestinal physiology Gut on a chip All pages with titles beginning with Gastrointestinal All pages with titles containing GastrointestinalReferences edit Overview of Invertebrates www ck12 org 6 October 2015 Retrieved 25 June 2021 Ruppert EE Fox RS Barnes RD 2004 Introduction to Bilateria Invertebrate Zoology 7 ed Brooks Cole p 197 1 ISBN 978 0 03 025982 1 gastrointestinal tract at Dorland s Medical Dictionary Gastrointestinal tract at the U S National Library of Medicine Medical Subject Headings MeSH digestive system at Dorland s Medical Dictionary G Hounnou C Destrieux J Desme P Bertrand S Velut 2002 12 01 Anatomical study of the length of the human intestine Surgical and Radiologic Anatomy 24 5 290 294 doi 10 1007 s00276 002 0057 y ISSN 0930 1038 PMID 12497219 S2CID 33366428 Raines Daniel Arbour Adrienne Thompson Hilary W Figueroa Bodine Jazmin Joseph Saju 2014 05 26 Variation in small bowel length Factor in achieving total enteroscopy Digestive Endoscopy 27 1 67 72 doi 10 1111 den 12309 ISSN 0915 5635 PMID 24861190 S2CID 19069407 Lin L Zhang J 2017 Role of intestinal microbiota and metabolites on gut homeostasis and human diseases BMC Immunology 18 1 2 doi 10 1186 s12865 016 0187 3 PMC 5219689 PMID 28061847 Marchesi J R Adams D H Fava F Hermes G D Hirschfield G M Hold G Quraishi M N Kinross J Smidt H Tuohy K M Thomas L V Zoetendal E G Hart A 2015 The gut microbiota and host health A new clinical frontier Gut 65 2 330 339 doi 10 1136 gutjnl 2015 309990 PMC 4752653 PMID 26338727 Clarke Gerard Stilling Roman M Kennedy Paul J Stanton Catherine Cryan John F Dinan Timothy G 2014 Minireview Gut Microbiota The Neglected Endocrine Organ Molecular Endocrinology 28 8 1221 38 doi 10 1210 me 2014 1108 PMC 5414803 PMID 24892638 Nelson RJ 2005 Introduction to Behavioral Endocrinology Sinauer Associates Massachusetts p 57 Thomasino Anne Marie 2001 Length of a Human Intestine The Physics Factbook Upper Gastrointestinal Tract at the U S National Library of Medicine Medical Subject Headings MeSH David A Warrell 2005 Oxford textbook of medicine Sections 18 33 Oxford University Press pp 511 ISBN 978 0 19 856978 7 Retrieved 1 July 2010 Lower Gastrointestinal Tract at the U S National Library of Medicine Medical Subject Headings MeSH Kapoor Vinay Kumar 13 Jul 2011 Gest Thomas R ed Large Intestine Anatomy Medscape WebMD LLC Retrieved 2013 08 20 Gray Henry 1918 Gray s Anatomy Philadelphia Lea amp Febiger a b Drake Richard L Vogl Wayne Tibbitts Adam W M Mitchell illustrations by Richard Richardson Paul 2015 Gray s anatomy for students 3rd ed Philadelphia Elsevier Churchill Livingstone p 312 ISBN 978 0 8089 2306 0 a b c Helander Herbert F Fandriks Lars 2014 06 01 Surface area of the digestive tract revisited Scandinavian Journal of Gastroenterology 49 6 681 689 doi 10 3109 00365521 2014 898326 ISSN 1502 7708 PMID 24694282 S2CID 11094705 Hall John 2011 Guyton and Hall textbook of medical physiology Twelfth ed Saunders Elsevier p 794 ISBN 9781416045748 Bruce M Carlson 2004 Human Embryology and Developmental Biology 3rd ed Saint Louis Mosby ISBN 978 0 323 03649 8 Abraham L Kierszenbaum 2002 Histology and cell biology an introduction to pathology St Louis Mosby ISBN 978 0 323 01639 1 a b c Sarna S K 2010 Introduction Colonic Motility From Bench Side to Bedside San Rafael California Morgan amp Claypool Life Sciences ISBN 9781615041503 The human proteome in gastrointestinal tract The Human Protein Atlas www proteinatlas org Retrieved 2017 09 21 Uhlen Mathias Fagerberg Linn Hallstrom Bjorn M Lindskog Cecilia Oksvold Per Mardinoglu Adil Sivertsson Asa Kampf Caroline Sjostedt Evelina 2015 01 23 Tissue based map of the human proteome Science 347 6220 1260419 doi 10 1126 science 1260419 ISSN 0036 8075 PMID 25613900 S2CID 802377 Gremel Gabriela Wanders Alkwin Cedernaes Jonathan Fagerberg Linn Hallstrom Bjorn Edlund Karolina Sjostedt Evelina Uhlen Mathias Ponten Fredrik 2015 01 01 The human gastrointestinal tract specific transcriptome and proteome as defined by RNA sequencing and antibody based profiling Journal of Gastroenterology 50 1 46 57 doi 10 1007 s00535 014 0958 7 ISSN 0944 1174 PMID 24789573 S2CID 21302849 a b Bowen Richard Gastrointestinal Transit How Long Does It Take Colorado State University Keendjele Tuwilika P T Eelu Hilja H Nashihanga Tunelago E Rennie Timothy W Hunter Christian John 1 March 2021 Corn When did I eat corn Gastrointestinal transit time in health science students Advances in Physiology Education 45 1 103 108 doi 10 1152 advan 00192 2020 PMID 33544037 S2CID 231817664 Kim SK 1968 Small intestine transit time in the normal small bowel study American Journal of Roentgenology 104 3 522 524 doi 10 2214 ajr 104 3 522 PMID 5687899 Ghoshal U C Sengar V Srivastava D 2012 Colonic Transit Study Technique and Interpretation Can These be Uniform Globally in Different Populations with Non uniform Colon Transit Time Journal of Neurogastroenterology and Motility 18 2 227 228 doi 10 5056 jnm 2012 18 2 227 PMC 3325313 PMID 22523737 Mowat Allan M Agace William W 2014 10 01 Regional specialization within the intestinal immune system Nature Reviews Immunology 14 10 667 685 doi 10 1038 nri3738 ISSN 1474 1741 PMID 25234148 S2CID 31460146 Flannigan Kyle L Geem Duke Harusato Akihito Denning Timothy L 2015 07 01 Intestinal Antigen Presenting Cells Key Regulators of Immune Homeostasis and Inflammation The American Journal of Pathology 185 7 1809 1819 doi 10 1016 j ajpath 2015 02 024 ISSN 1525 2191 PMC 4483458 PMID 25976247 Sanchez de Medina Fermin Romero Calvo Isabel Mascaraque Cristina Martinez Augustin Olga 2014 12 01 Intestinal inflammation and mucosal barrier function Inflammatory Bowel Diseases 20 12 2394 2404 doi 10 1097 MIB 0000000000000204 ISSN 1536 4844 PMID 25222662 S2CID 11434730 Schubert Mitchell L 2014 11 01 Gastric secretion Current Opinion in Gastroenterology 30 6 578 582 doi 10 1097 MOG 0000000000000125 ISSN 1531 7056 PMID 25211241 S2CID 8267813 Marquez Mercedes Fernandez Gutierrez Del Alamo Clotilde Giron Gonzalez Jose Antonio 2016 01 28 Gut epithelial barrier dysfunction in human immunodeficiency virus hepatitis C virus coinfected patients Influence on innate and acquired immunity World Journal of Gastroenterology 22 4 1433 1448 doi 10 3748 wjg v22 i4 1433 ISSN 2219 2840 PMC 4721978 PMID 26819512 Furusawa Yukihiro Obata Yuuki Fukuda Shinji Endo Takaho A Nakato Gaku Takahashi Daisuke Nakanishi Yumiko Uetake Chikako Kato Keiko Kato Tamotsu Takahashi Masumi Fukuda Noriko N Murakami Shinnosuke Miyauchi Eiji Hino Shingo Atarashi Koji Onawa Satoshi Fujimura Yumiko Lockett Trevor Clarke Julie M Topping David L Tomita Masaru Hori Shohei Ohara Osamu Morita Tatsuya Koseki Haruhiko Kikuchi Jun Honda Kenya Hase Koji Ohno Hiroshi 2013 Commensal microbe derived butyrate induces the differentiation of colonic regulatory T cells Nature 504 7480 446 450 Bibcode 2013Natur 504 446F doi 10 1038 nature12721 PMID 24226770 S2CID 4408815 Knight Judson 2002 Science of Everyday Things Real life biology Vol 4 Gale ISBN 9780787656348 Jakoby WB Ziegler DM 5 December 1990 The enzymes of detoxication The Journal of Biological Chemistry 265 34 20715 8 doi 10 1016 S0021 9258 17 45272 0 PMID 2249981 Joel Lucas 10 January 2020 Fossil Reveals Earth s Oldest Known Animal Guts The find in a Nevada desert revealed an intestine inside a creature that looks like a worm made of a stack of ice cream cones The New York Times Retrieved 10 January 2020 Svhiffbauer James D et al 10 January 2020 Discovery of bilaterian type through guts in cloudinomorphs from the terminal Ediacaran Period Nature Communications 11 205 205 Bibcode 2020NatCo 11 205S doi 10 1038 s41467 019 13882 z PMC 6954273 PMID 31924764 Nielsen C Brunet T amp Arendt D Evolution of the bilaterian mouth and anus Nat Ecol Evol 2 1358 1376 2018 https doi org 10 1038 s41559 018 0641 0 De Robertis E M amp Tejeda Munoz N 2022 Evo Devo of urbilateria and its larval forms Developmental Biology 487 10 20 https doi org 10 1016 j ydbio 2022 04 003 Nitzan Orna Elias Mazen Peretz Avi Saliba Walid 2016 01 21 Role of antibiotics for treatment of inflammatory bowel disease World Journal of Gastroenterology 22 3 1078 1087 doi 10 3748 wjg v22 i3 1078 ISSN 1007 9327 PMC 4716021 PMID 26811648 Wier LM Steiner CA Owens PL February 2015 Surgeries in Hospital Owned Outpatient Facilities 2012 HCUP Statistical Brief 188 Rockville MD Agency for Healthcare Research and Quality Fox James Timothy Wang January 2007 Inflammation Atrophy and Gastric Cancer Journal of Clinical Investigation review 117 1 60 69 doi 10 1172 JCI30111 PMC 1716216 PMID 17200707 Murphy Kenneth 20 May 2014 Janeway s Immunobiology New York Garland Science Taylor and Francis Group LLC pp 389 398 ISBN 978 0 8153 4243 4 Parham Peter 20 May 2014 The Immune System New York Garland Science Taylor and Francis Group LLC p 494 ISBN 978 0 8153 4146 8 Goering Richard 20 May 2014 MIMS Medical Microbiology Philadelphia Elsevier pp 32 64 294 133 4 208 303 4 502 ISBN 978 0 3230 4475 2 Hiskey Daven 12 November 2010 Violin strings were never made out of actual cat guts TodayIFoundOut com Retrieved 15 December 2015 World s oldest condom Ananova 2008 Retrieved 2008 04 11 External links edit nbsp Look up gastrointestinal tract gastrointestinal or tract in Wiktionary the free dictionary nbsp Wikimedia Commons has media related to wbr Gastrointestinal tract and wbr Digestive system nbsp The Wikibook Human Physiology has a page on the topic of The gastrointestinal system The gastro intestinal tract in the Human Protein Atlas Your Digestive System and How It Works at National Institutes of Health Retrieved from https en wikipedia org w index php title Gastrointestinal tract amp oldid 1206582010, wikipedia, wiki, book, books, library,

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