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Gastrointestinal physiology is the branch of human physiology that addresses the physical function of the gastrointestinal (GI) tract. The function of the GI tract is to process ingested food by mechanical and chemical means, extract nutrients and excrete waste products. The GI tract is composed of the alimentary canal, that runs from the mouth to the anus, as well as the associated glands, chemicals, hormones, and enzymes that assist in digestion. The major processes that occur in the GI tract are: motility, secretion, regulation, digestion and circulation. The proper function and coordination of these processes are vital for maintaining good health by providing for the effective digestion and uptake of nutrients.


Motility

The gastrointestinal tract generates
motility Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
using
smooth muscle Smooth muscle is an involuntary non- striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit ...
subunits linked by gap junctions. These subunits fire spontaneously in either a tonic or a phasic fashion. Tonic contractions are those contractions that are maintained from several minutes up to hours at a time. These occur in the sphincters of the tract, as well as in the anterior stomach. The other type of contractions, called phasic contractions, consist of brief periods of both relaxation and contraction, occurring in the posterior stomach and the small intestine, and are carried out by the muscularis externa. Motility may be overactive (hypermotility), leading to diarrhea or vomiting, or underactive (hypomotility), leading to constipation or vomiting; either may cause abdominal pain.


Stimulation

The stimulation for these contractions likely originates in modified smooth muscle cells called interstitial cells of Cajal. These cells cause spontaneous cycles of
slow wave potential A slow-wave potential is a rhythmic electrophysiological event in the gastrointestinal tract. The normal conduction of slow waves is one of the key regulators of gastrointestinal motility. Slow waves are generated and propagated by a class of pac ...
s that can cause
action potentials An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
in smooth muscle cells. They are associated with the contractile smooth muscle via gap junctions. These slow wave potentials must reach a threshold level for the action potential to occur, whereupon Ca2+ channels on the smooth muscle open and an action potential occurs. As the contraction is graded based upon how much Ca2+ enters the cell, the longer the duration of slow wave, the more action potentials occur. This, in turn, results in greater contraction force from the smooth muscle. Both amplitude and duration of the slow waves can be modified based upon the presence of
neurotransmitters A neurotransmitter is a signaling molecule secreted by a neuron to affect another cell across a synapse. The cell receiving the signal, any main body part or target cell, may be another neuron, but could also be a gland or muscle cell. Neuro ...
, hormones or other paracrine signaling. The number of slow wave potentials per minute varies based upon the location in the digestive tract. This number ranges from 3 waves/min in the stomach to 12 waves/min in the intestines.


Contraction patterns

The patterns of GI contraction as a whole can be divided into two distinct patterns,
peristalsis Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, which ...
and segmentation. Occurring between meals, the migrating motor complex is a series of peristaltic wave cycles in distinct phases starting with relaxation, followed by an increasing level of activity to a peak level of peristaltic activity lasting for 5–15 minutes. This cycle repeats every 1.5–2 hours but is interrupted by food ingestion. The role of this process is likely to clean excess bacteria and food from the digestive system.


Peristalsis

Peristalsis is one of the patterns that occur during and shortly after a meal. The contractions occur in wave patterns traveling down short lengths of the GI tract from one section to the next. The contractions occur directly behind the
bolus Bolus may refer to: Geography * Bolus, Iran, a village in Ardabil Province, Iran * Bolus, or Baulus, an Anatolian village on the site of ancient Berissa Medicine * Bolus (digestion), a ball-shaped mass moving through the digestive tract * Bolus ...
of food that is in the system, forcing it toward the anus into the next relaxed section of smooth muscle. This relaxed section then contracts, generating smooth forward movement of the bolus at between 2–25 cm per second. This contraction pattern depends upon hormones, paracrine signals, and the autonomic nervous system for proper regulation.


Segmentation

Segmentation also occurs during and shortly after a meal within short lengths in segmented or random patterns along the intestine. This process is carried out by the longitudinal muscles relaxing while circular muscles contract at alternating sections thereby mixing the food. This mixing allows food and digestive enzymes to maintain a uniform composition, as well as to ensure contact with the epithelium for proper absorption.


Secretion

Every day, seven liters of fluid are secreted by the digestive system. This fluid is composed of four primary components: ions, digestive enzymes, mucus, and bile. About half of these fluids are secreted by the salivary glands, pancreas, and liver, which compose the accessory organs and glands of the digestive system. The rest of the fluid is secreted by the GI epithelial cells.


Ions

The largest component of secreted fluids is ions and water, which are first secreted and then reabsorbed along the tract. The ions secreted primarily consist of H+, K+, Cl, HCO3 and Na+. Water follows the movement of these ions. The GI tract accomplishes this ion pumping using a system of proteins that are capable of
active transport In cellular biology, ''active transport'' is the movement of molecules or ions across a cell membrane from a region of lower concentration to a region of higher concentration—against the concentration gradient. Active transport requires cellu ...
, facilitated diffusion and open channel ion movement. The arrangement of these proteins on the apical and basolateral sides of the epithelium determines the net movement of ions and water in the tract. H+ and Cl are secreted by the parietal cells into the lumen of the stomach creating acidic conditions with a low pH of 1. H+ is pumped into the stomach by exchanging it with K+. This process also requires ATP as a source of energy; however, Cl then follows the positive charge in the H+ through an open apical channel protein. HCO3 secretion occurs to neutralize the acid secretions that make their way into the
duodenum The duodenum is the first section of the small intestine in most higher vertebrates, including mammals, reptiles, and birds. In fish, the divisions of the small intestine are not as clear, and the terms anterior intestine or proximal intestine ...
of the small intestine. Most of the HCO3 comes from pancreatic acinar cells in the form of NaHCO3 in an aqueous solution. This is the result of the high concentration of both HCO3 and Na+ present in the duct creating an osmotic gradient to which the water follows.


Digestive enzymes

The second vital secretion of the GI tract is that of digestive enzymes that are secreted in the mouth, stomach and intestines. Some of these enzymes are secreted by accessory digestive organs, while others are secreted by the epithelial cells of the stomach and intestine. While some of these enzymes remain embedded in the wall of the GI tract, others are secreted in an inactive proenzyme form. When these proenzymes reach the lumen of the tract, a factor specific to a particular proenzyme will activate it. A prime example of this is
pepsin Pepsin is an endopeptidase that breaks down proteins into smaller peptides. It is produced in the gastric chief cells of the stomach lining and is one of the main digestive enzymes in the digestive systems of humans and many other animals, ...
, which is secreted in the stomach by chief cells. Pepsin in its secreted form is inactive (
pepsinogen Pepsin is an endopeptidase that breaks down proteins into smaller peptides. It is produced in the gastric chief cells of the stomach lining and is one of the main digestive enzymes in the digestive systems of humans and many other animals, w ...
). However, once it reaches the gastric lumen it becomes activated into
pepsin Pepsin is an endopeptidase that breaks down proteins into smaller peptides. It is produced in the gastric chief cells of the stomach lining and is one of the main digestive enzymes in the digestive systems of humans and many other animals, ...
by the high H+ concentration, becoming an enzyme vital to digestion. The release of the enzymes is regulated by neural, hormonal, or paracrine signals. However, in general, parasympathetic stimulation increases secretion of all digestive enzymes.


Mucus

Mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
is released in the stomach and intestine, and serves to lubricate and protect the inner mucosa of the tract. It is composed of a specific family of
glycoproteins Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glyco ...
termed mucins and is generally very viscous. Mucus is made by two types of specialized cells termed mucus cells in the stomach and goblet cells in the intestines. Signals for increased mucus release include parasympathetic innervations, immune system response and enteric nervous system messengers.


Bile

Bile is secreted into the duodenum of the small intestine via the
common bile duct The common bile duct, sometimes abbreviated as CBD, is a duct in the gastrointestinal tract of organisms that have a gallbladder. It is formed by the confluence of the common hepatic duct and cystic duct and terminates by uniting with pancr ...
. It is produced in liver cells and stored in the gall bladder until release during a meal. Bile is formed of three elements: bile salts,
bilirubin Bilirubin (BR) (Latin for "red bile") is a red-orange compound that occurs in the normal catabolic pathway that breaks down heme in vertebrates. This catabolism is a necessary process in the body's clearance of waste products that arise from the ...
and cholesterol. Bilirubin is a waste product of the breakdown of hemoglobin. The cholesterol present is secreted with the feces. The bile salt component is an active non-enzymatic substance that facilitates fat absorption by helping it to form an emulsion with water due to its
amphoteric In chemistry, an amphoteric compound () is a molecule or ion that can react both as an acid and as a base. What exactly this can mean depends on which definitions of acids and bases are being used. One type of amphoteric species are amphip ...
nature. These salts are formed in the hepatocytes from bile acids combined with an
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
. Other compounds such as the waste products of drug degradation are also present in the bile.


Regulation

The digestive system has a complex system of motility and secretion regulation which is vital for proper function. This task is accomplished via a system of long reflexes from the
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
(CNS), short reflexes from the
enteric nervous system The enteric nervous system (ENS) or intrinsic nervous system is one of the main divisions of the autonomic nervous system (ANS) and consists of a mesh-like system of neurons that governs the function of the gastrointestinal tract. It is capable o ...
(ENS) and reflexes from GI
peptides Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. ...
working in harmony with each other.


Long reflexes

Long reflexes to the digestive system involve a sensory neuron sending information to the brain, which integrates the signal and then sends messages to the digestive system. While in some situations, the sensory information comes from the GI tract itself; in others, information is received from sources other than the GI tract. When the latter situation occurs, these reflexes are called feedforward reflexes. This type of reflex includes reactions to food or danger triggering effects in the GI tract. Emotional responses can also trigger GI response such as the butterflies in the stomach feeling when nervous. The feedforward and emotional reflexes of the GI tract are considered cephalic reflexes.


Short reflexes

Control of the digestive system is also maintained by ENS, which can be thought of as a digestive brain that can help to regulate motility, secretion and growth. Sensory information from the digestive system can be received, integrated and acted upon by the enteric system alone. When this occurs, the reflex is called a short reflex. Although this may be the case in several situations, the ENS can also work in conjunction with the CNS; vagal afferents from the viscera are received by the medulla, efferents are affected by the
vagus nerve The vagus nerve, also known as the tenth cranial nerve, cranial nerve X, or simply CN X, is a cranial nerve that interfaces with the parasympathetic control of the heart, lungs, and digestive tract. It comprises two nerves—the left and rig ...
. When this occurs, the reflex is called vagovagal reflex. The
myenteric plexus The myenteric plexus (or Auerbach's plexus) provides motor innervation to both layers of the muscular layer of the gut, having both parasympathetic and sympathetic input (although present ganglion cell bodies belong to parasympathetic innervation ...
and submucosal plexus are both located in the gut wall and receive sensory signals from the lumen of the gut or the CNS.


Gastrointestinal peptides

''For further information see Gastrointestinal hormone'' GI peptides are signal molecules that are released into the blood by the GI cells themselves. They act on a variety of tissues including the brain, digestive accessory organs, and the GI tract. The effects range from excitatory or inhibitory effects on motility and secretion to feelings of satiety or hunger when acting on the brain. These hormones fall into three major categories, the gastrin and
secretin Secretin is a hormone that regulates water homeostasis throughout the body and influences the environment of the duodenum by regulating secretions in the stomach, pancreas, and liver. It is a peptide hormone produced in the S cells of the duoden ...
families, with the third composed of all the other hormones unlike those in the other two families. Further information on the GI peptides is summarized in the table below.


Digestion

*
carbohydrates In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or may ...
(
monosaccharide Monosaccharides (from Greek '' monos'': single, '' sacchar'': sugar), also called simple sugars, are the simplest forms of sugar and the most basic units (monomers) from which all carbohydrates are built. They are usually colorless, water-sol ...
,
disaccharide A disaccharide (also called a double sugar or ''biose'') is the sugar formed when two monosaccharides are joined by glycosidic linkage. Like monosaccharides, disaccharides are simple sugars soluble in water. Three common examples are sucrose, ...
) *
proteins Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respondi ...
* lipids


Splanchnic circulation

*
Superior mesenteric artery In human anatomy, the superior mesenteric artery (SMA) is an artery which arises from the anterior surface of the abdominal aorta, just inferior to the origin of the celiac trunk, and supplies blood to the intestine from the lower part of the ...
*
Inferior mesenteric artery In human anatomy, the inferior mesenteric artery, often abbreviated as IMA, is the third main branch of the abdominal aorta and arises at the level of L3, supplying the large intestine from the distal transverse colon to the upper part of the an ...


External links


Notes
at
University of Bristol The University of Bristol is a Red brick university, red brick Russell Group research university in Bristol, England. It received its royal charter in 1909, although it can trace its roots to a Society of Merchant Venturers, Merchant Venturers' sc ...
*


Notes and references

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