The enteric nervous system (ENS) is one of the three divisions of the

autonomic nervous system The autonomic nervous system (ANS), sometimes called the visceral nervous system and formerly the vegetative nervous system, is a division of the nervous system that operates viscera, internal organs, smooth muscle and glands. The autonomic nervo ...

(ANS), the others being the

sympathetic nervous system The sympathetic nervous system (SNS or SANS, sympathetic autonomic nervous system, to differentiate it from the somatic nervous system) is one of the three divisions of the autonomic nervous system, the others being the parasympathetic nervous sy ...

(SNS) and

parasympathetic nervous system The parasympathetic nervous system (PSNS) is one of the three divisions of the autonomic nervous system, the others being the sympathetic nervous system and the enteric nervous system. The autonomic nervous system is responsible for regulat ...

(PSNS). It consists of a mesh-like system of

neuron A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...

s that governs the function of the

gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the Digestion, digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascula ...

. The ENS is nicknamed the "second brain". It is derived from

neural crest The neural crest is a ridge-like structure that is formed transiently between the epidermal ectoderm and neural plate during vertebrate development. Neural crest cells originate from this structure through the epithelial-mesenchymal transition, ...

cells. The enteric nervous system is capable of operating independently of the brain and spinal cord, but is thought to rely on innervation from the

vagus nerve The vagus nerve, also known as the tenth cranial nerve (CN X), plays a crucial role in the autonomic nervous system, which is responsible for regulating involuntary functions within the human body. This nerve carries both sensory and motor fibe ...

and prevertebral ganglia in healthy subjects. However, studies have shown that the system is operable with a severed vagus nerve. The neurons of the enteric nervous system control the motor functions of the system, in addition to the secretion of gastrointestinal enzymes. These neurons communicate through many neurotransmitters similar to the CNS, including

acetylcholine Acetylcholine (ACh) is an organic compound that functions in the brain and body of many types of animals (including humans) as a neurotransmitter. Its name is derived from its chemical structure: it is an ester of acetic acid and choline. Par ...

dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic chemical of the catecholamine and phenethylamine families. It is an amine synthesized ...

, and

serotonin Serotonin (), also known as 5-hydroxytryptamine (5-HT), is a monoamine neurotransmitter with a wide range of functions in both the central nervous system (CNS) and also peripheral tissues. It is involved in mood, cognition, reward, learning, ...

. The large presence of serotonin and dopamine in the

intestines 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 tract is the largest of the body's systems, after the cardiovascular system. ...

are key areas of research for neurogastroenterology.

Structure

2402 Layers of the Gastrointestinal Tract

The enteric nervous system in humans consists of some 500 million

s (including the various types of Dogiel cells), 0.5% of the number of neurons in the

brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...

, five times as many as the one hundred million neurons in the human spinal cord, and about as many as in the whole nervous system of a cat. The enteric nervous system is embedded in the lining of the

gastrointestinal system 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 tract is the largest of the body's systems, after the cardiovascular system. T ...

, beginning in the

esophagus The esophagus (American English), oesophagus (British English), or œsophagus (Œ, archaic spelling) (American and British English spelling differences#ae and oe, see spelling difference) all ; : ((o)e)(œ)sophagi or ((o)e)(œ)sophaguses), c ...

and extending down to the anus. The neurons of the ENS are collected into two types of

ganglia A ganglion (: ganglia) is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system, this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system, there a ...

: myenteric (Auerbach's) and submucosal (Meissner's) plexuses. Myenteric plexuses are located between the inner and outer layers of the muscularis externa, while submucosal plexuses are located in the

submucosa The submucosa (or tela submucosa) is a thin layer of tissue in various organs of the gastrointestinal, respiratory, and genitourinary tracts. It is the layer of dense irregular connective tissue that supports the mucosa (mucous membrane) an ...

Auerbach's plexus

Auerbach's plexus, also known as the myenteric plexus, is a collection of fibers and postganglionic autonomic cell bodies that lie between the circular and longitudinal layers of the muscularis externa in the gastrointestinal tract. It was discovered and named by German neuropathologist Leopold Auerbach. These neurons provide motor inputs to both layers of the muscularis externa and provide both parasympathetic and sympathetic input. The anatomy of the plexus is similar to the anatomy of the

central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain, spinal cord and retina. The CNS is so named because the brain integrates the received information and coordinates and influences the activity o ...

. The plexus includes sensory receptors, such as

chemoreceptors A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance ( endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorece ...

and mechanoreceptors, that are used to provide sensory input to the interneurons in the enteric nervous system. The plexus is the parasympathetic nucleus of origin for the vagus nerve and communicates with the

medulla oblongata The medulla oblongata or simply medulla is a long stem-like structure which makes up the lower part of the brainstem. It is anterior and partially inferior to the cerebellum. It is a cone-shaped neuronal mass responsible for autonomic (involun ...

through both the anterior and posterior vagal nerves.

Submucosal plexus

The submucosal plexus (also known as Meissner's plexus) is found in the submucosal layer of the

. It was discovered and named by German physiologist Georg Meissner. It functions as a pathway for the innervation in the mucosa layer of the gastrointestinal wall.

Function

The ENS is capable of autonomous functions like the coordination of

reflex In biology, a reflex, or reflex action, is an involuntary, unplanned sequence or action and nearly instantaneous response to a stimulus. Reflexes are found with varying levels of complexity in organisms with a nervous system. A reflex occurs ...

es; although it receives considerable innervation from the autonomic nervous system, it can and does operate independently of the brain and the spinal cord. Its study is the focus of neurogastroenterology.

Complexity

The enteric nervous system is a complex part of the nervous system. The enteric nervous system can operate autonomously. It normally communicates with the

(CNS) through the

parasympathetic The parasympathetic nervous system (PSNS) is one of the three divisions of the autonomic nervous system, the others being the sympathetic nervous system and the enteric nervous system. The autonomic nervous system is responsible for regulat ...

(e.g., via the

) and sympathetic (e.g., via the prevertebral ganglia) nervous systems. However,

vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...

studies show that when the

is severed, the enteric nervous system continues to function. In vertebrates, the enteric nervous system includes

efferent neurons Efferent nerve fibers are axons (nerve fibers) of efferent neurons that exit a particular region. These terms have a slightly different meaning in the context of the peripheral nervous system (PNS) and central nervous system (CNS). The efferent ...

, afferent neurons, and

interneurons Interneurons (also called internuncial neurons, association neurons, connector neurons, or intermediate neurons) are neurons that are not specifically motor neurons or sensory neurons. Interneurons are the central nodes of neural circuits, ena ...

, all of which make the enteric nervous system capable of carrying reflexes and acting as an integrating center in the absence of CNS input. The sensory neurons report on mechanical and chemical conditions. Through intestinal muscles, the motor neurons control

peristalsis Peristalsis ( , ) is a type of intestinal motility, characterized by symmetry in biology#Radial symmetry, radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an wikt:anterograde, anterograde dir ...

and churning of intestinal contents. Other neurons control the secretion of

enzymes An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as pro ...

. The enteric nervous system also makes use of more than 30 neurotransmitters, most of which are identical to the ones found in CNS, such as

, and

. More than 90% of the body's serotonin lies in the gut, as well as about 50% of the body's dopamine, which is currently being studied to further our understanding of its utility in the brain. The enteric nervous system has the capacity to alter its response depending on such factors as bulk and nutrient composition. In addition, the ENS contains support cells which are similar to astroglia of the brain and a diffusion barrier around the capillaries surrounding ganglia which is similar to the

blood–brain barrier The blood–brain barrier (BBB) is a highly selective semipermeable membrane, semipermeable border of endothelium, endothelial cells that regulates the transfer of solutes and chemicals between the circulatory system and the central nervous system ...

of cerebral blood vessels.

Peristalsis

Peristalsis is a series of radially symmetrical contractions and relaxations of muscles which propagate down a muscular tube. In humans and other mammals, peristalsis is found in the smooth muscles of the digestive tract to propel contents through the digestive system. The word is derived from New Latin and comes from the Greek peristallein, "to wrap around," from peri-, "around" + stellein, "draw in, bring together; set in order". Peristalsis was discovered in 1899 by the work of physiologists

William Bayliss Sir William Maddock Bayliss (2 May 1860 – 27 August 1924) was an English physiologist. Life He was born in Wednesbury, Staffordshire but shortly thereafter his father, a successful merchant of ornamental ironwork, moved his family to a ...

and

Ernest Starling Ernest Henry Starling (17 April 1866 – 2 May 1927) was a British physiologist who contributed many fundamental ideas to this subject. These ideas were important parts of the British contribution to physiology, which at that time led the world. ...

. Working on the

small intestines The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through the ...

of dogs, they found that the response of increasing the pressure in the intestine caused the contraction of the muscle wall above the point of stimulation and the relaxation of the muscle wall below the point of stimulation.

Segmentation

Segmentation contractions are the contractions in intestines carried out by the smooth muscle walls. Unlike peristalsis, which involves the contraction and relaxation of muscles in one direction, segmentation occurs simultaneously in both directions as the circular muscles alternatively contract. This allows for thorough mixing of intestinal contents, known as

chyme Chyme or chymus (; ) is the semi-fluid mass of partly digested food that is expelled by the stomach, through the pyloric valve, into the duodenum (the beginning of the small intestine). Chyme results from the mechanical and chemical breakdown ...

, to allow greater absorption.

Secretion

The secretion of gastrointestinal hormones, such as

gastrin Gastrin is a peptide hormone that stimulates secretion of gastric acid (HCl) by the parietal cells of the stomach and aids in gastric motility. It is released by G cells in the pyloric antrum of the stomach, duodenum, and the pancreas. ...

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 duodenum ...

, is regulated through cholinergic neurons residing in the walls of the digestive tract. Hormone secretion is controlled by the vagovagal reflex, where the neurons in the digestive tract communicate through both afferent and efferent pathways with the

Clinical significance

Neurogastroenterology encompasses the study of the brain, the gut, and their interactions with relevance to the understanding and management of gastrointestinal

motility Motility is the ability of an organism to move independently using metabolism, metabolic energy. This biological concept encompasses movement at various levels, from whole organisms to cells and subcellular components. Motility is observed in ...

and functional gastrointestinal disorders. Specifically, neurogastroenterology focuses on the functions, malfunctions, and the malformations of the sympathetic,

, and enteric divisions of the digestive tract. The term also describes a medical sub-specialism of gastroenterology dedicated to the treatment of motility and functional gastrointestinal disorders.

Functional gastrointestinal disorders

Functional gastrointestinal (GI) disorders are a class of gastrointestinal disorders where there is a malfunction in the normal activities of the gastrointestinal tract, but there are no structural abnormalities that can explain the cause. There are rarely any tests that can detect the presence of these disorders. Clinical research in neurogastroenterology focuses mainly on the study of common functional gastrointestinal disorders such as

irritable bowel syndrome Irritable bowel syndrome (IBS) is a functional gastrointestinal disorder characterized by a group of symptoms that commonly include abdominal pain, abdominal bloating, and changes in the consistency of bowel movements. These symptoms may ...

, the most common functional GI disorder.

Motility disorders

Motility disorders are the second classification of gastrointestinal disorder studied by neurogastroenterologists. Motility disorders are divided by what they affect, with four regions: The esophagus, the stomach, the small intestines, and the large intestines. Clinical research in neurogastroenterology focuses mainly on the study of common motility disorders such as

gastroesophageal reflux disease Gastroesophageal reflux disease (GERD) or gastro-oesophageal reflux disease (GORD) is a chronic upper gastrointestinal disease in which stomach content persistently and regularly flows up into the esophagus, resulting in symptoms and/or ...

, the damage of the mucosa of the esophagus caused by rising stomach acid through the lower esophageal sphincter.

Gut ischemia

ENS function can be damaged by

ischemia Ischemia or ischaemia is a restriction in blood supply to any tissue, muscle group, or organ of the body, causing a shortage of oxygen that is needed for cellular metabolism (to keep tissue alive). Ischemia is generally caused by problems ...

. Transplantation, previously described as a theoretical possibility, has been a clinical reality in the United States since 2011 and is regularly performed at some hospitals.

Additional images

Image:Gray1071.png, The myenteric plexus of a rabbit. X 50. Image:Gray1072.png, The submucosal plexus of a rabbit. X 50.

Neurogastroenterology societies

* American Neurogastroenterology and Motility Society * European Society of Neurogastroenterology and MotilityESNM - European Society for Neurogastroenterology & Motility
/ref> ** '' Neurogastroenterology & Motility''

References

Further references

* * * {{DEFAULTSORT:Enteric Nervous System Animal anatomy