Stomach

The stomach is a muscular, hollow organ in the gastrointestinal

Stomach
tract of humans and many other animals, including several
invertebrates. The stomach has a dilated structure and functions
as a vital organ in the digestive system. The stomach is
involved in the gastric phase of digestion, following chewing.
It performs a chemical breakdown by means of enzymes and
hydrochloric acid.

In humans and many other animals, the stomach is located

between the esophagus and the small intestine. The stomach
secretes digestive enzymes and gastric acid to aid in food
digestion. The pyloric sphincter controls the passage of
partially digested food (chyme) from the stomach into the
duodenum, where peristalsis takes over to move this through
the rest of the intestines.

Structure
In the human digestive system, the stomach lies between the
esophagus and the duodenum (the first part of the small
intestine). It is in the left upper quadrant of the abdominal Scheme of digestive tract, with
cavity. The top of the stomach lies against the diaphragm. stomach in red
Lying behind the stomach is the pancreas. A large double fold
of visceral peritoneum called the greater omentum hangs down
from the greater curvature of the stomach. Two sphincters keep
the contents of the stomach contained; the lower esophageal
sphincter (found in the cardiac region), at the junction of the
esophagus and stomach, and the pyloric sphincter at the
junction of the stomach with the duodenum.

The stomach is surrounded by parasympathetic (stimulant) and

sympathetic (inhibitor) plexuses (networks of blood vessels and
nerves in the anterior gastric, posterior, superior and inferior, Sections of the human stomach
celiac and myenteric), which regulate both the secretory
activity of the stomach and the motor (motion) activity of its Details
muscles. Precursor Foregut
System Digestive system
Because it is a distensible organ, it normally expands to hold
about one litre of food.[3] The stomach of a newborn human Artery Right gastric artery, left
baby will only be able to retain about 30 millilitres. The gastric artery, right gastro-
maximum stomach volume in adults is between 2 and 4 omental artery, left gastro-
litres.[4][5] omental artery, short
gastric arteries

Sections Vein Right gastric vein, left

gastric vein, right
In classical anatomy the human stomach is divided into four gastroepiploic vein, left
sections, beginning at the cardia.[7] gastroepiploic vein, short
gastric veins
The cardia is where the contents of the esophagus
Nerve Celiac ganglia, vagus
empty into the stomach.[8]
nerve[1]
The fundus (from Latin 'bottom') is formed in the
upper curved part. Lymph Celiac lymph nodes[2]
The body is the main, central region of the stomach. Identifiers
The pylorus (from Greek 'gatekeeper') is the lower Latin ventriculus
section of the stomach that empties contents into the
duodenum. Greek gaster
MeSH D013270 (https://meshb.nl
The cardia is defined as the region following the "z-line" of the m.nih.gov/record/ui?ui=D01
gastroesophageal junction, the point at which the epithelium
3270)
changes from stratified squamous to columnar. Near the cardia
is the lower esophageal sphincter.[8] Research has shown that TA98 A05.5.01.001 (https://ifaa.u
the cardia is not an anatomically distinct region of the stomach nifr.ch/Public/EntryPage/TA
but a region of the esophageal lining damaged by reflux.[9] 98%20Tree/Entity%20TA9
8%20EN/05.5.01.001%20E
ntity%20TA98%20EN.htm)
Anatomical proximity
TA2 2901 (https://ta2viewer.ope
The stomach bed refers to the structures upon which the nanatomy.org/?id=2901)
stomach rests in mammals.[10][11] These include the tail of the FMA 7148 (https://bioportal.bioo
pancreas, splenic artery, left kidney, left suprarenal gland, ntology.org/ontologies/FM
transverse colon and its mesocolon, and the left crus of A/?p=classes&conceptid=ht
diaphragm, and the left colic flexure. The term was introduced tp%3A%2F%2Fpurl.org%2
Fsig%2Font%2Ffma%2Ffm
a7148)
Anatomical terminology

1. Body of stomach 2. Fundus 3.

Anterior wall 4. Greater curvature 5.
Lesser curvature 6. Cardia 9. Pyloric
sphincter 10. Pyloric antrum 11.
Pyloric canal 12. Angular incisure 13.
Gastric canal 14. Rugae[6]
around 1896 by Philip Polson of the Catholic University School of Medicine, Dublin. However this was
brought into disrepute by surgeon anatomist J Massey.[12][13][14]

Blood supply

Schematic image of the blood supply to the human stomach: left and right gastric artery, left and right
gastroepiploic artery and short gastric artery[15]

The lesser curvature of the human stomach is supplied by the right gastric artery inferiorly and the left
gastric artery superiorly, which also supplies the cardiac region. The greater curvature is supplied by the
right gastroepiploic artery inferiorly and the left gastroepiploic artery superiorly. The fundus of the stomach,
and also the upper portion of the greater curvature, is supplied by the short gastric arteries, which arise from
the splenic artery.

Microanatomy

Wall

Like the other parts of the gastrointestinal tract, the human stomach
walls consist of a mucosa, submucosa, muscularis externa,
subserosa and serosa.[16]

The inner part of the lining of the stomach, the gastric mucosa,
consists of an outer layer of column-shaped cells, a lamina propria,
and a thin layer of smooth muscle called the muscularis mucosa. The gastrointestinal wall of the
Beneath the mucosa lies the submucosa, consisting of fibrous human stomach
connective tissue.[17] Meissner's plexus is in this layer interior to
the oblique muscle layer.[18]

Outside of the submucosa lies another muscular layer, the muscularis externa. It consists of three layers of
muscular fibres, with fibres lying at angles to each other. These are the inner oblique, middle circular, and
outer longitudinal layers. The presence of the inner oblique layer is distinct from other parts of the
gastrointestinal tract, which do not possess this layer.[19] Stomach contains the thickest muscularis layer
consisting of three layers, thus maximum peristalsis occurs here.

The inner oblique layer: This layer is responsible for creating the motion that churns and
physically breaks down the food. It is the only layer of the three which is not seen in other
parts of the digestive system. The antrum has thicker skin cells in its walls and performs
more forceful contractions than the fundus.
 The middle circular layer: At this layer, the pylorus is surrounded by a thick circular muscular
wall, which is normally tonically constricted, forming a functional (if not anatomically
discrete) pyloric sphincter, which controls the movement of chyme into the duodenum. This
layer is concentric to the longitudinal axis of the stomach.
Auerbach's plexus (myenteric plexus) is found between the outer longitudinal and the
middle circular layer and is responsible for the innervation of both (causing peristalsis and
mixing).

The outer longitudinal layer is responsible for moving the bolus towards the pylorus of the stomach
through muscular shortening.

To the outside of the muscularis externa lies a serosa, consisting of layers of connective tissue continuous
with the peritoneum.

Glands

The mucosa lining the stomach is lined with a number of these pits,
which receive gastric juice, secreted by between 2 and 7 gastric
glands. Gastric juice is an acidic fluid containing hydrochloric acid
and the digestive enzyme pepsin. The glands contains a number of
cells, with the function of the glands changing depending on their
position within the stomach.

Within the body and fundus of the stomach lie the fundic glands. In
general, these glands are lined by column-shaped cells that secrete a
Histology of normal fundic mucosa.
protective layer of mucus and bicarbonate. Additional cells present
Fundic glands are simple, branched
include parietal cells that secrete hydrochloric acid and intrinsic
tubular glands that extend from the
factor, chief cells that secrete pepsinogen (this is a precursor to
bottom of the gastric pits to the
pepsin- the highly acidic environment converts the pepsinogen to
muscularis mucosae; the more
pepsin), and neuroendocrine cells that secrete serotonin.[20] distinctive cells are parietal cells.
H&E stain.
Glands differ where the stomach meets the esophagus and near the
pylorus.[21] Near the junction between the stomach and the
esophagus lie cardiac glands, which primarily secrete mucus.[20]
They are fewer in number than the other gastric glands and are
more shallowly positioned in the mucosa. There are two kinds -
either simple tubular with short ducts or compound racemose
resembling the duodenal Brunner's glands. Near the pylorus lie
pyloric glands located in the antrum of the pylorus. They secrete
mucus, as well as gastrin produced by their G cells.[22]

Gene and protein expression Histology of normal antral mucosa.

Antral mucosa is formed by
About 20,000 protein coding genes are expressed in human cells branched coiled tubular glands lined
and nearly 70% of these genes are expressed in the normal by secretory cells similar in
stomach.[23][24] Just over 150 of these genes are more specifically appearance to the surface mucus
expressed in the stomach compared to other organs, with only some cells. H&E stain.
20 genes being highly specific. The corresponding specific proteins
expressed in stomach are mainly involved in creating a suitable
environment for handling the digestion of food for uptake of nutrients. Highly stomach-specific proteins
include GKN1, expressed in the mucosa; pepsinogen PGC and the lipase LIPF, expressed in chief cells;
and gastric ATPase ATP4A and gastric intrinsic factor GIF, expressed in parietal cells.[25]

Development
In early human embryogenesis, the ventral part of the embryo abuts the yolk sac. During the third week of
development, as the embryo grows, it begins to surround parts of the sac. The enveloped portions form the
basis for the adult gastrointestinal tract.[26] The sac is surrounded by a network of vitelline arteries and
veins. Over time, these arteries consolidate into the three main arteries that supply the developing
gastrointestinal tract: the celiac artery, superior mesenteric artery, and inferior mesenteric artery. The areas
supplied by these arteries are used to define the foregut, midgut, and hindgut.[26] The surrounded sac
becomes the primitive gut. Sections of this gut begin to differentiate into the organs of the gastrointestinal
tract, and the esophagus, and stomach form from the foregut.[26]

Function

Digestion

In the human digestive system, a bolus (a small rounded mass of chewed up food) enters the stomach
through the esophagus via the lower esophageal sphincter. The stomach releases proteases (protein-
digesting enzymes such as pepsin) and hydrochloric acid, which kills or inhibits bacteria and provides the
acidic pH of 2 for the proteases to work. Food is churned by the stomach through muscular contractions of
the wall called peristalsis – reducing the volume of the bolus, before looping around the fundus[27] and the
body of stomach as the boluses are converted into chyme (partially digested food). Chyme slowly passes
through the pyloric sphincter and into the duodenum of the small intestine, where the extraction of nutrients
begins.

Gastric juice in the stomach also contains pepsinogen. Hydrochloric acid activates this inactive form of
enzyme into the active form, pepsin. Pepsin breaks down proteins into polypeptides.

Mechanical digestion

Within a few moments after food enters the stomach, mixing waves begin to occur at intervals of
approximately 20 seconds. A mixing wave is a unique type of peristalsis that mixes and softens the food
with gastric juices to create chyme. The initial mixing waves are relatively gentle, but these are followed by
more intense waves, starting at the body of the stomach and increasing in force as they reach the pylorus.

The pylorus, which holds around 30 mL of chyme, acts as a filter, permitting only liquids and small food
particles to pass through the mostly, but not fully, closed pyloric sphincter. In a process called gastric
emptying, rhythmic mixing waves force about 3 mL of chyme at a time through the pyloric sphincter and
into the duodenum. Release of a greater amount of chyme at one time would overwhelm the capacity of the
small intestine to handle it. The rest of the chyme is pushed back into the body of the stomach, where it
continues mixing. This process is repeated when the next mixing waves force more chyme into the
duodenum.
Gastric emptying is regulated by both the stomach and the duodenum. The presence of chyme in the
duodenum activates receptors that inhibit gastric secretion. This prevents additional chyme from being
released by the stomach before the duodenum is ready to process it.[28]

Chemical digestion

The fundus stores both undigested food and gases that are released during the process of chemical
digestion. Food may sit in the fundus of the stomach for a while before being mixed with the chyme. While
the food is in the fundus, the digestive activities of salivary amylase continue until the food begins mixing
with the acidic chyme. Ultimately, mixing waves incorporate this food with the chyme, the acidity of which
inactivates salivary amylase and activates lingual lipase. Lingual lipase then begins breaking down
triglycerides into free fatty acids, and mono- and diglycerides.

The breakdown of protein begins in the stomach through the actions of HCl and the enzyme pepsin.

The contents of the stomach are completely emptied into the duodenum within 2 to 4 hours after the meal is
eaten. Different types of food take different amounts of time to process. Foods heavy in carbohydrates
empty fastest, followed by high-protein foods. Meals with a high triglyceride content remain in the stomach
the longest. Since enzymes in the small intestine digest fats slowly, food can stay in the stomach for 6 hours
or longer when the duodenum is processing fatty chyme. However, this is still a fraction of the 24 to 72
hours that full digestion typically takes from start to finish.[28]

Absorption

Although the absorption in the human digestive system is mainly a function of the small intestine, some
absorption of certain small molecules nevertheless does occur in the stomach through its lining. This
includes:

Water, if the body is dehydrated

Medication, such as aspirin
Amino acids[29]
10–20% of ingested ethanol (e.g. from alcoholic beverages)[30]
Caffeine[31]
To a small extent water-soluble vitamins (most are absorbed in the small intestine)[32]
The parietal cells of the human stomach are responsible for producing intrinsic factor, which is necessary
for the absorption of vitamin B12. B12 is used in cellular metabolism and is necessary for the production of
red blood cells, and the functioning of the nervous system.

Control of secretion and motility

Emptying of stomach chyme into the duodenum through the pyloric sphincter

Chyme from the stomach is slowly released into the duodenum through coordinated peristalsis and opening
of the pyloric sphincter. The movement and the flow of chemicals into the stomach are controlled by both
the autonomic nervous system and by the various digestive hormones of the digestive system:

The hormone gastrin causes an increase in the secretion of HCl from the parietal cells and
pepsinogen from chief cells in the stomach. It also causes increased motility in the stomach.
Gastrin Gastrin is released by G cells in the stomach in response to distension of the antrum and
digestive products (especially large quantities of incompletely digested proteins). It is
inhibited by a pH normally less than 4(high acid), as well as the hormone somatostatin.
Cholecystokinin (CCK) has most effect on the gall bladder, causing gall bladder contractions,
but it also decreases gastric emptying and increases release of pancreatic juice, which is
Cholecystokinin
alkaline and neutralizes the chyme. CCK is synthesized by I-cells in the mucosal epithelium
of the small intestine.

In a different and rare manner, secretin, which has the most effects on the pancreas, also
Secretin diminishes acid secretion in the stomach. Secretin is synthesized by S-cells, which are
located in the duodenal mucosa as well as in the jejunal mucosa in smaller numbers.

Gastric inhibitory Gastric inhibitory peptide (GIP) decreases both gastric acid release and motility. GIP is
peptide synthesized by K-cells, which are located in the duodenal and jejunal mucosa.
Enteroglucagon Enteroglucagon decreases both gastric acid and motility.

Other than gastrin, these hormones all act to turn off the stomach action. This is in response to food
products in the liver and gall bladder, which have not yet been absorbed. The stomach needs to push food
into the small intestine only when the intestine is not busy. While the intestine is full and still digesting food,
the stomach acts as storage for food.

Other

Effects of EGF

Epidermal growth factor (EGF) results in cellular proliferation, differentiation, and survival.[33] EGF is a
low-molecular-weight polypeptide first purified from the mouse submandibular gland, but since then found
in many human tissues including the submandibular gland, and the parotid gland. Salivary EGF, which also
seems to be regulated by dietary inorganic iodine, also plays an important physiological role in the
maintenance of oro-esophageal and gastric tissue integrity. The biological effects of salivary EGF include
healing of oral and gastroesophageal ulcers, inhibition of gastric acid secretion, stimulation of DNA
synthesis, and mucosal protection from intraluminal injurious factors such as gastric acid, bile acids, pepsin,
and trypsin and from physical, chemical, and bacterial agents.[34]

Stomach as nutrition sensor

The human stomach has receptors responsive to sodium glutamate[35] and this information is passed to the
lateral hypothalamus and limbic system in the brain as a palatability signal through the vagus nerve.[36] The
stomach can also sense, independently of tongue and oral taste receptors, glucose,[37] carbohydrates,[38]
proteins,[38] and fats.[39] This allows the brain to link nutritional value of foods to their tastes.[37]

Thyrogastric syndrome

This syndrome defines the association between thyroid disease and chronic gastritis, which was first
described in the 1960s.[40] This term was coined also to indicate the presence of thyroid autoantibodies or
autoimmune thyroid disease in patients with pernicious anemia, a late clinical stage of atrophic gastritis.[41]
In 1993, a more complete investigation on the stomach and thyroid was published,[42] reporting that the
thyroid is, embryogenetically and phylogenetically, derived from a primitive stomach, and that the thyroid
cells, such as primitive gastroenteric cells, migrated and specialized in uptake of iodide and in storage and
elaboration of iodine compounds during vertebrate evolution. In fact, the stomach and thyroid share iodine-
concentrating ability and many morphological and functional similarities, such as cell polarity and apical
microvilli, similar organ-specific antigens and associated autoimmune diseases, secretion of glycoproteins
(thyroglobulin and mucin) and peptide hormones, the digesting and readsorbing ability, and lastly, similar
ability to form iodotyrosines by peroxidase activity, where iodide acts as an electron donor in the presence
of H2 O2 . In the following years, many researchers published reviews about this syndrome.[43]

Clinical significance

Diseases

A series of radiographs can be used to examine the stomach for

various disorders. This will often include the use of a barium
swallow. Another method of examination of the stomach, is the use
of an endoscope. A gastric emptying study is considered the gold
standard to assess the gastric emptying rate.[44]

A large number of studies have indicated that most cases of peptic An endoscopy of a normal stomach
ulcers, and gastritis, in humans are caused by Helicobacter pylori of a healthy 65-year-old woman
infection, and an association has been seen with the development of
stomach cancer.[45]

A stomach rumble is actually noise from the intestines.

Surgery

In humans, many bariatric surgery procedures involve the stomach, in order to lose weight. A gastric band
may be placed around the cardia area, which can adjust to limit intake. The anatomy of the stomach may be
modified, or the stomach may be bypassed entirely.
Surgical removal of the stomach is called a gastrectomy, and
removal of the cardia area is a called a cardiectomy.
"Cardiectomy" is a term that is also used to describe the removal of
the heart.[46][47][48] A gastrectomy may be carried out because of
gastric cancer or severe perforation of the stomach wall.

Fundoplication is stomach surgery in which the fundus is wrapped

around the lower esophagus and stitched into place. It is used to
treat gastroesophageal reflux disease (GERD).[49]

History
Endoscopic image of a fundic gland
polyp
There were previously conflicting statements in the academic
anatomy community[50][51][52] over whether the cardia is part of
the stomach, part of the esophagus or a distinct entity. Modern
surgical and medical textbooks have agreed that "the gastric cardia is now clearly considered to be part of
the stomach."[53][54]

Etymology

The word stomach is derived from Greek stomachos (στόμαχος), ultimately from stoma (στόμα)
'mouth'.[55] Gastro- and gastric (meaning 'related to the stomach') are both derived from Greek gaster
(γαστήρ) 'belly'.[56][57][58]

Other animals
Although the precise shape and size of the stomach varies widely among different vertebrates, the relative
positions of the esophageal and duodenal openings remain relatively constant. As a result, the organ always
curves somewhat to the left before curving back to meet the pyloric sphincter. However, lampreys,
hagfishes, chimaeras, lungfishes, and some teleost fish have no stomach at all, with the esophagus opening
directly into the intestine. These animals all consume diets that require little storage of food, no predigestion
with gastric juices, or both.[59]

Yellow
Esophagus
Green
Esophageal (nonglandular)
region.[63]
Comparison of stomach glandular regions from several Purple
mammalian species. Frequency of glands may vary Cardiac gland region.[63]
more smoothly between regions than is diagrammed Red
here. Asterisk (ruminant) represents the omasum, which Fundic gland region.[63]
is absent in Tylopoda (Tylopoda also have some cardiac Blue
glands opening onto ventral reticulum and rumen[60]) Pyloric gland region.[63]
Many other variations exist among the mammals.[61][62] Dark blue
Duodenum
The gastric lining is usually divided into two regions, an anterior portion lined by fundic glands and a
posterior portion lined with pyloric glands. Cardiac glands are unique to mammals, and even then are
absent in a number of species. The distributions of these glands vary between species, and do not always
correspond with the same regions as in humans. Furthermore, in many non-human mammals, a portion of
the stomach anterior to the cardiac glands is lined with epithelium essentially identical to that of the
esophagus. Ruminants, in particular, have a complex stomach, the first three chambers of which are all
lined with esophageal mucosa.[59]

In birds and crocodilians, the stomach is divided into two regions. Anteriorly is a narrow tubular region, the
proventriculus, lined by fundic glands, and connecting the true stomach to the crop. Beyond lies the
powerful muscular gizzard, lined by pyloric glands, and, in some species, containing stones that the animal
swallows to help grind up food.[59]

In insects there is also a crop. The insect stomach is called the midgut.
Information about the stomach in echinoderms or molluscs can be found under the respective articles.

Additional images

Greater omentum A more realistic An autopsy of a Human stomach

and stomach of image, showing the human stomach,
humans celiac artery and its showing the many
branches in folds (rugae) of the
humans; the liver stomach. 2012
has been raised, Instituto Nacional de
and the lesser Cardiología.
omentum and
anterior layer of the
greater omentum
removed.

High-quality image
of the stomach

See also
Gastroesophageal reflux disease
Human gastrointestinal microbiota
Proton-pump inhibitor

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External links
Stomach (https://www.proteinatlas.org/humanproteome/stomach) at the Human Protein Atlas
Digestion of proteins in the stomach or tiyan (https://web.archive.org/web/20070310213747/
http://www.med.uiuc.edu/m1/biochemistry/TA%20reviews/sam/AminoAcids.htm) (archived
10 March 2007)
Site with details of how ruminants process food (https://web.archive.org/web/200910270338
03/http://uk.geocities.com/bacterial_ed/bacteria_and_food.htm) (archived 27 October 2009)
Control of Gastric Emptying (http://www.partone.lifeinthefastlane.com/control_of_gastric_em
ptying.html) (Archived (https://web.archive.org/web/20191112051823/http://www.partone.lifei
nthefastlane.com/control_of_gastric_emptying.html) 2019-11-12 at the Wayback Machine)

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