| Gastrointestinal tract | |
|---|---|
 Diagram of the gastrointestinal tract in the averagehuman | |
| Details | |
| System | Digestive system |
| Identifiers | |
| Latin | tractus digestorius (mouth toanus), canalis alimentarius (esophagus tolarge intestine), canalis gastrointestinales (stomach tolarge intestine) |
| MeSH | D041981 |
| Anatomical terminology | |
 |
| Major parts of the |
| Gastrointestinal tract |
|---|
Thegastrointestinal tract (GI tract,digestive tract,alimentary canal) is the tract or passageway of thedigestive system that leads from themouth to theanus. The GI tract contains all the majororgans of the digestive system, in humans and other animals, including theesophagus,stomach, andintestines. Food taken in through the mouth isdigested to extractnutrients and absorbenergy, and the waste expelled at the anus asfeces.Gastrointestinal is an adjective meaning of or pertaining to the stomach and intestines.
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, whilecnidarians andacoels have a single pore for both digestion and excretion.[1][2]
The human gastrointestinal tract consists of theesophagus, stomach, and intestines, and is divided into the upper and lower gastrointestinal tracts.[3] The GI tract includes all structures between themouth and theanus,[4] forming a continuous passageway that includes the main organs of digestion, namely, thestomach,small intestine, andlarge intestine. The completehuman digestive system is made up of the gastrointestinal tract plus the accessory organs of digestion (thetongue,salivary glands,pancreas,liver andgallbladder).[5] The tract may also be divided intoforegut,midgut, andhindgut, reflecting theembryological origin of each segment. The whole human GI tract is about nine meters (30 feet) long atautopsy. It is considerably shorter in the living body because the intestines, which are tubes ofsmooth muscle tissue, maintain constantmuscle tone in a halfway-tense state but can relax in spots to allow for local distention andperistalsis.[6][7]
The gastrointestinal tract contains thegut microbiota, with some 1,000 differentstrains ofbacteria having diverse roles in the maintenance ofimmune health andmetabolism, and many othermicroorganisms.[8][9][10] Cells of the GI tract releasehormones to help regulate the digestive process. Thesedigestive hormones, includinggastrin,secretin,cholecystokinin, andghrelin, are mediated through eitherintracrine orautocrine mechanisms, indicating that the cells releasing these hormones are conserved structures throughoutevolution.[11]
The structure and function can be described both asgross anatomy and asmicroscopic anatomy orhistology. The tract itself is divided into upper and lower tracts, and the intestinessmall andlarge parts.[12]
The upper gastrointestinal tract consists of themouth,pharynx,esophagus,stomach, andduodenum.[13]The exact demarcation between the upper and lower tracts is thesuspensory muscle of the duodenum. This differentiates the embryonic borders between the foregut and midgut, and is also the division commonly used by clinicians to describegastrointestinal bleeding as being of either "upper" or "lower" origin. Upondissection, 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 thejejunum.
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 theembryonicmesoderm.
The lower gastrointestinal tract includes most of thesmall intestine and all of thelarge intestine.[15] Inhuman anatomy, theintestine (bowel orgut; Greek:éntera) is the segment of the gastrointestinal tract extending from the pyloric sphincter of thestomach to theanus and as in other mammals, consists of two segments: thesmall intestine and thelarge intestine. In humans, the small intestine is further subdivided into theduodenum,jejunum, andileum while the large intestine is subdivided into thececum, ascending, transverse, descending, and sigmoidcolon,rectum, andanal canal.[16][17]
Thesmall intestine begins at theduodenum and is a tubular structure, usually between 6 and 7 m long.[18] Itsmucosal area in an adult human is about 30 m2 (320 sq ft).[19] The combination of thecircular 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:
Thelarge intestine forms an arch starting at thececum and ending at therectum andanal canal. It also includes theappendix, which is attached to thececum. 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] The longest part of the large intestine is thecolon whose main function is to absorb water and salts.[21]
The large intestine begins at the cecum, where theappendix is located. This is also the start of the colon as theascending colon in the back wall of the abdomen. At theright colic flexure (hepatic flexure) (the flexed portion of the ascending andtransverse colon) it runs across the abdomen in the transverse colon, passing below the diaphragm. At theleft colic flexure (splenic flexure) the flexed portion of the transverse anddescending colon, it descends down the left side of the abdomen.It reaches thesigmoid colon which is a loop of the colon closest to the rectum and continues to the rectum andanal canal.
The gut is anendoderm-derived structure. At approximately the sixteenth day of human development, theembryo begins to foldventrally (with the embryo's ventral surface becomingconcave) 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 theyolk sac, anendoderm-lined structure in contact with theventral aspect of the embryo, begins to be pinched off to become the primitive gut. The yolk sac remains connected to the gut tube via thevitelline duct. Usually, this structure regresses during development; in cases where it does not, it is known asMeckel's diverticulum.
Duringfetal life, the primitive gut is gradually patterned into three segments:foregut,midgut, andhindgut. 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 thestomach andcolon, 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.[22]
| 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 theceliac 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 | lowerduodenum,jejunum,ileum,cecum,appendix,ascending colon, and first two-thirds of thetransverse colon | branches of thesuperior mesenteric artery |
| Hindgut | last third of the transverse colon, to the upper part of the anal canal | last third of thetransverse colon,descending colon,rectum, and upper part of theanal canal | branches of theinferior mesenteric artery |
The gastrointestinal tract has a form of general histology with some differences that reflect the specialization in functional anatomy.[23] The GI tract can be divided into four concentric layers in the following order:
The mucosa is the innermost layer of the gastrointestinal tract. The mucosa surrounds thelumen, or open space within the tube. This layer comes in direct contact with digested food (chyme). The mucosa is made up of:
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.
The submucosa consists of a dense irregular layer of connective tissue with large blood vessels, lymphatics, and nerves branching into the mucosa andmuscularis externa. It contains thesubmucosal plexus, anenteric nervous plexus, situated on the inner surface of themuscularis externa.
Themuscular layer consists of an inner circular layer and alongitudinal 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.[24] 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 the outer longitudinal layer.
Between the circular and longitudinal muscle layers is themyenteric plexus. This controls peristalsis. Activity is initiated by the pacemaker cells, (myentericinterstitial 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 theautonomic nervous system.[24]
The coordinated contractions of these layers is calledperistalsis 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 aschyme. In the large intestine, the remaining semi-solid substance is referred to asfaeces.[24]
The outermost layer of the gastrointestinal tract consists of several layers ofconnective tissue.
Intraperitoneal parts of the GI tract are covered withserosa. These include most of thestomach, first part of theduodenum, all of thesmall intestine,caecum andappendix,transverse colon,sigmoid colon andrectum. In these sections of the gut, there is a clear boundary between the gut and the surrounding tissue. These parts of the tract have amesentery.
Retroperitoneal parts are covered withadventitia. They blend into the surrounding tissue and are fixed in position. For example, the retroperitoneal section of the duodenum usually passes through thetranspyloric plane. These include theesophagus,pylorus of the stomach, distalduodenum,ascending colon,descending colon andanal canal. In addition, theoral cavity has adventitia.
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.[25][26] 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 arepepsinogen PGC and thelipase LIPF, expressed inchief cells, and gastricATPase ATP4A andgastric intrinsic factor GIF, expressed inparietal cells of the stomach mucosa. Specific proteins expressed in the stomach and duodenum involved in defence includemucin proteins, such asmucin 6 andintelectin-1.[27]
The time taken for food to transit through the gastrointestinal tract varies on multiple factors, including age, ethnicity, and gender.[28][29] Several techniques have been used to measure transit time, including radiography following abarium-labeled meal, breathhydrogen analysis,scintigraphic analysis following aradiolabeled meal,[30] and simple ingestion and spotting ofcorn kernels.[31] 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.[32] Total emptying of the stomach takes around 4–5 hours, and transit through the colon takes 30 to 50 hours.[30][33][34]
The gastrointestinal tract forms an important part of theimmune system.[35]
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 theexposed surface of the skin), these immune components function to prevent pathogens from entering the blood and lymph circulatory systems.[36] Fundamental components of this protection are provided by theintestinal mucosal barrier, which is composed of physical, biochemical, and immune elements elaborated by the intestinal mucosa.[37] Microorganisms also are kept at bay by an extensive immune system comprising thegut-associated lymphoid tissue (GALT)
There are additional factors contributing to protection from pathogen invasion. For example, lowpH (ranging from 1 to 4) of the stomach is fatal for manymicroorganisms that enter it.[38] Similarly,mucus (containingIgAantibodies) neutralizes many pathogenic microorganisms.[39] Other factors in the GI tract contribution to immune function includeenzymes secreted in thesaliva andbile.
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.[40] It has been demonstrated that the intake of a high fiber diet could be responsible for the induction ofT-regulatory cells (Tregs). This is due to the production ofshort-chain fatty acids during the fermentation of plant-derived nutrients such asbutyrate andpropionate. Basically, the butyrate induces the differentiation of Treg cells by enhancinghistone H3acetylation in the promoter and conserved non-coding sequence regions of theFOXP3 locus, thus regulating theT cells, resulting in the reduction of the inflammatory response and allergies.
The large intestine contains multiple types ofbacteria that can break down molecules the human body cannot process alone,[41] demonstrating asymbiotic relationship. These bacteria are responsible for gas production athost–pathogen interface, which is released asflatulence. Intestinal bacteria can also participate in biosynthesis reactions. For example, certain strains in the large intestine produce vitamin B12;[42] an essential compound in humans for things like DNA synthesis and red blood cell production.[43] However, the primary function of the large intestine is water absorption from digested material (regulated by thehypothalamus) and the reabsorption ofsodium and nutrients.[44]
Beneficialintestinal bacteria compete with potentially harmfulbacteria 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 maintaininghomeostasis.[citation needed] An imbalanced ratio results indysbiosis.
Enzymes such asCYP3A4, along with theantiporter activities, are also instrumental in the intestine's role ofdrug metabolism in the detoxification ofantigens andxenobiotics.[45]
In mostvertebrates, includingamphibians,birds,reptiles,egg-laying mammals, and somefish, the gastrointestinal tract ends in acloaca and not ananus. In the cloaca, theurinary 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 subgroupPlacentalia have even separate urinary and genital openings.
Duringearly development, the asymmetric position of the bowels and inner organs is initiated (see alsoaxial twist theory).
Ruminants show many specializations for digesting andfermenting tough plant material, consisting ofadditional stomach compartments.
Many birds and other animals have a specialised stomach in the digestive tract called agizzard used for grinding up food.
Another feature found in a range of animals is thecrop. 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 theCloudinidae was discovered; it lived during the lateEdiacaranperiod about 550 million years ago.[46][47]
A through-gut (one with both mouth and anus) is thought to have evolved within thenephrozoan clade ofBilateria, after their ancestral ventral orifice (single, as incnidarians andacoels; re-evolved in nephrozoans likeflatworms) 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 plusgenital opening). A stretched gut without the middle part closed is present in another branch of bilaterians, the extinctproarticulates. This and theamphistomic 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.[48][49]
There are many diseases and conditions that can affect the gastrointestinal system, includinginfections,inflammation andcancer.
Variouspathogens, such asbacteria that causefoodborne illnesses, can inducegastroenteritis which results frominflammation of the stomach and small intestine.Antibiotics to treat such bacterial infections can decrease themicrobiome diversity of the gastrointestinal tract, and further enable inflammatory mediators.[50] 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 asdiverticulitis.
Inflammatory bowel disease is an inflammatory condition affecting the bowel walls, and includes the subtypesCrohn's disease andulcerative 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 anautoimmune 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 isirritable bowel syndrome. Functional constipation andchronic functional abdominal pain are other functional disorders of the intestine that have physiological causes but do not have identifiable structural, chemical, or infectious pathologies.
Several symptoms can indicate problems with the gastrointestinal tract, including:
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.[52]
Various methods of imaging the gastrointestinal tract include theupper andlower gastrointestinal series:
Intestines from animals other than humans are used in a number of ways. From each species oflivestock that is a source ofmilk, a correspondingrennet is obtained from the intestines of milk-fedcalves.Pig andcalf intestines are eaten, and pig intestines are used assausage casings. Calf intestines supplycalf-intestinal alkaline phosphatase (CIP), and are used to makegoldbeater's skin.Other uses are:
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