Sunday, 13 March 2011


The nervous system derives its name from nerves, which are cylindrical bundles of fibers that emanate from the brain and central cord, and branch repeatedly to innervate every part of the body.[2] Nerves are large enough to have been recognized by the ancient Egyptians, Greeks, and Romans,[3] but their internal structure was not understood until it became possible to examine them using a microscope.[4] A microscopic examination shows that nerves consist primarily of the axons of neurons, along with a variety of membranes that wrap around them and segregate them into fascicles. The neurons that give rise to nerves do not lie entirely within the nerves themselves—their cell bodies reside within the brain, central cord, or peripheral ganglia.[2]

All animals more advanced than sponges have nervous systems. However, even sponges, unicellular animals, and non-animals such as slime molds have cell-to-cell signalling mechanisms that are precursors to those of neurons.[5] In radially symmetric animals such as the jellyfish and hydra, the nervous system consists of a diffuse network of isolated cells.[6] In bilaterian animals, which make up the great majority of existing species, the nervous system has a common structure that originated early in the Cambrian period, over 500 million years ago.[7]

The nervous system is primarily made up of two categories of cells: neurons and glial cells.
Structure of a typical neuron Neuron
At one end of an elongated structure is a branching mass. At the centre of this mass is the nucleus and the branches are dendrites. A thick axon trails away from the mass, ending with further branching which are labeled as axon terminals. Along the axon are a number of protuberances labeled as myelin sheaths.
Node of
Axon terminal
Schwann cell
Myelin sheath

The nervous system is defined by the presence of a special type of cell—the neuron (sometimes called "neurone" or "nerve cell").[2] Neurons can be distinguished from other cells in a number of ways, but their most fundamental property is that they communicate with other cells via synapses, which are membrane-to-membrane junctions containing molecular machinery that allows rapid transmission of signals, either electrical or chemical.[2] Many types of neuron possess an axon, a protoplasmic protrusion that can extend to distant parts of the body and make thousands of synaptic contacts.[8] Axons frequently travel through the body in bundles called nerves.

Even in the nervous system of a single species such as humans, hundreds of different types of neurons exist, with a wide variety of morphologies and functions.[8] These include sensory neurons that transmute physical stimuli such as light and sound into neural signals, and motor neurons that transmute neural signals into activation of muscles or glands; however in many species the great majority of neurons receive all of their input from other neurons and send their output to other neurons.[2]
 Glial cells

Glial cells are non-neuronal cells that provide support and nutrition, maintain homeostasis, form myelin, and participate in signal transmission in the nervous system.[9] In the human brain, it is estimated that the total number of glia roughly equals the number of neurons, although the proportions vary in different brain areas.[10] Among the most important functions of glial cells are to support neurons and hold them in place; to supply nutrients to neurons; to insulate neurons electrically; to destroy pathogens and remove dead neurons; and to provide guidance cues directing the axons of neurons to their targets.[9] A very important type of glial cell (oligodendrocytes in the central nervous system, and Schwann cells in the peripheral nervous system) generates layers of a fatty substance called myelin that wraps around axons and provides electrical insulation which allows them to transmit action potentials much more rapidly and efficiently.
 Anatomy in vertebrates
Wiki letter w cropped.svg     This section requires expansion.
Diagram showing the major divisions of the vertebrate nervous system.

The nervous system of vertebrate animals (including humans) is divided into the central nervous system (CNS) and peripheral nervous system (PNS).[11]

The central nervous system (CNS) is the largest part, and includes the brain and spinal cord.[11] The spinal cavity contains the spinal cord, while the head contains the brain. The CNS is enclosed and protected by meninges, a three-layered system of membranes, including a tough, leathery outer layer called the dura mater. The brain is also protected by the skull, and the spinal cord by the vertebrae.

The peripheral nervous system (PNS) is a collective term for the nervous system structures that do not lie within the CNS.[12] The large majority of the axon bundles called nerves are considered to belong to the PNS, even when the cell bodies of the neurons to which they belong reside within the brain or spinal cord. The PNS is divided into somatic and visceral parts. The somatic part consists of the nerves that innervate the skin, joints, and muscles. The cell bodies of somatic sensory neurons lie in dorsal root ganglia of the spinal cord. The visceral part, also known as the autonomic nervous system, contains neurons that innervate the internal organs, blood vessels, and glands. The autonomic nervous system itself consists of two parts: the sympathetic nervous system and the parasympathetic nervous system. Some authors also include sensory neurons whose cell bodies lie in the periphery (for senses such as hearing) as part of the PNS; others, however, omit them.[13]
Horizontal bisection of the head of an adult man, showing skin, skull, and brain with grey matter (brown in this image) and underlying white matter

The vertebrate nervous system can also be divided into areas called grey matter ("gray matter" in American spelling) and white matter.[14] Grey matter (which is only grey in preserved tissue, and is better described as pink or light brown in living tissue) contains a high proportion of cell bodies of neurons. White matter is composed mainly of myelinated axons, and takes its color from the myelin. White matter includes all of the peripheral nerves, and much of the interior of the brain and spinal cord. Grey matter is found in clusters of neurons in the brain and spinal cord, and in cortical layers that line their surfaces. There is an anatomical convention that a cluster of neurons in the brain or spinal cord is called a nucleus, whereas a cluster of neurons in the periphery is called a ganglion.[15] There are, however, a few exceptions to this rule, notably including the part of the forebrain called the basal ganglia.[16]
 Comparative anatomy and evolution
 Neural precursors in sponges

Sponges have no cells connected to each other by synaptic junctions, that is, no neurons, and therefore no nervous system. They do, however, have homologs of many genes that play key roles in synaptic function. Recent studies have shown that sponge cells express a group of proteins that cluster together to form a structure resembling a postsynaptic density (the signal-receiving part of a synapse).[5] However, the function of this structure is currently unclear. Although sponge cells do not show synaptic transmission, they do communicate with each other via calcium waves and other impulses, which mediate some simple actions such as whole-body contraction.[17]

Jellyfish, comb jellies, and related animals have diffuse nerve nets rather than a central nervous system. In most jellyfish the nerve net is spread more or less evenly across the body; in comb jellies it is concentrated near the mouth. The nerve nets consist of sensory neurons that pick up chemical, tactile, and visual signals, motor neurons that can activate contractions of the body wall, and intermediate neurons that detect patterns of activity in the sensory neurons and send signals to groups of motor neurons as a result. In some cases groups of intermediate neurons are clustered into discrete ganglia.[6]

The development of the nervous system in radiata is relatively unstructured. Unlike bilaterians, radiata only have two primordial cell layers, endoderm and ectoderm. Neurons are generated from a special set of ectodermal precursor cells, which also serve as precursors for every other ectodermal cell type.[18]
A rod-shaped body contains a digestive system running from the mouth at one end to the anus at the other. Alongside the digestive system is a nerve cord with a brain at the end, near to the mouth.
Nervous system of a bilaterian animal, in the form of a nerve cord with segmental enlargements, and a "brain" at the front

The vast majority of existing animals are bilaterians, meaning animals with left and right sides that are approximate mirror images of each other. All bilateria are thought to have descended from a common wormlike ancestor that appeared in the Cambrian period, 550–600 million years ago.[7] The fundamental bilaterian body form is a tube with a hollow gut cavity running from mouth to anus, and a nerve cord with an enlargement (a "ganglion") for each body segment, with an especially large ganglion at the front, called the "brain".
Area of the human body surface innervated by each spinal nerve

Even mammals, including humans, show the segmented bilaterian body plan at the level of the nervous system. The spinal cord contains a series of segmental ganglia, each giving rise to motor and sensory nerves that innervate a portion of the body surface and underlying musculature. On the limbs, the layout of the innervation pattern is complex, but on the trunk it gives rise to a series of narrow bands. The top three segments belong to the brain, giving rise to the forebrain, midbrain, and hindbrain.[19]

Bilaterians can be divided, based on events that occur very early in embryonic development, into two groups (superphyla) called protostomes and deuterostomes.[20] Deuterostomes include vertebrates as well as echinoderms, hemichordates (mainly acorn worms), and Xenoturbellidans.[21] Protostomes, the more diverse group, include arthropods, molluscs, and numerous types of worms. There is a basic difference between the two groups in the placement of the nervous system within the body: protostomes possess a nerve cord on the ventral (usually bottom) side of the body, whereas in deuterostomes the nerve cord is on the dorsal (usually top) side. In fact, numerous aspects of the body are inverted between the two groups, including the expression patterns of several genes that show dorsal-to-ventral gradients. Most anatomists now consider that the bodies of protostomes and deuterostomes are "flipped over" with respect to each other, a hypothesis that was first proposed by Geoffroy Saint-Hilaire for insects in comparison to vertebrates. Thus insects, for example, have nerve cords that run along the ventral midline of the body, while all vertebrates have spinal cords that run along the dorsal midline.[22]
    This section's factual accuracy is disputed. Please see the relevant discussion on the talk page. (May 2010)
Earthworm nervous system. Top: side view of the front of the worm. Bottom: nervous system in isolation, viewed from above

Worms are the simplest bilaterian animals, and reveal the basic structure of the bilaterian nervous system in the most straightforward way. As an example, earthworms have dual nerve cords running along the length of the body and merging at the tail and the mouth. These nerve cords are connected by transverse nerves like the rungs of a ladder. These transverse nerves help coordinate the two sides of the animal. Two ganglia at the head end function similar to a simple brain. Photoreceptors on the animal's eyespots provide sensory information on light and dark.[23]

The nervous system of one very small worm, the roundworm Caenorhabditis elegans, has been mapped out down to the synaptic level. Every neuron and its cellular lineage has been recorded and most, if not all, of the neural connections are known. In this species, the nervous system is sexually dimorphic; the nervous systems of the two sexes, males and hermaphrodites, have different numbers of neurons and groups of neurons that perform sex-specific functions. In C. elegans, males have exactly 383 neurons, while hermaphrodites have exactly 302 neurons.[24]
Internal anatomy of a spider, showing the nervous system in blue

Arthropods, such as insects and crustaceans, have a nervous system made up of a series of ganglia, connected by a ventral nerve cord made up of two parallel connectives running along the length of the belly.[25] Typically, each body segment has one ganglion on each side, though some ganglia are fused to form the brain and other large ganglia. The head segment contains the brain, also known as the supraesophageal ganglion. In the insect nervous system, the brain is anatomically divided into the protocerebrum, deutocerebrum, and tritocerebrum. Immediately behind the brain is the subesophageal ganglion, which is composed of three pairs of fused ganglia. It controls the mouthparts, the salivary glands and certain muscles. Many arthropods have well-developed sensory organs, including compound eyes for vision and antennae for olfaction and pheromone sensation. The sensory information from these organs is processed by the brain.

In insects, many neurons have cell bodies that are positioned at the edge of the brain and are electrically passive—the cell bodies serve only to provide metabolic support and do not participate in signalling. A protoplasmic fiber runs from the cell body and branches profusely, with some parts transmitting signals and other parts receiving signals. Thus, most parts of the insect brain have passive cell bodies arranged around the periphery, while the neural signal processing takes place in a tangle of protoplasmic fibers called neuropil, in the interior.[26]
 "Identified" neurons

A neuron is called identified if it has properties that distinguish it from every other neuron in the same animal—properties such as location, neurotransmitter, gene expression pattern, and connectivity—and if every individual organism belonging to the same species has one and only one neuron with the same set of properties.[27] In vertebrate nervous systems very few neurons are "identified" in this sense—in humans, there are believed to be none—but in simpler nervous systems, some or all neurons may be thus unique. In the roundworm C. elegans, whose nervous system is the most thoroughly described of any animal's, every neuron in the body is uniquely identifiable, with the same location and the same connections in every individual worm. One notable consequence of this fact is that the form of the C. elegans nervous system is completely specified by the genome, with no experience-dependent plasticity.[24]

The brains of many molluscs and insects also contain substantial numbers of identified neurons.[27] In vertebrates, the best known identified neurons are the gigantic Mauthner cells of fish.[28] Every fish has two Mauthner cells, located in the bottom part of the brainstem, one on the left side and one on the right. Each Mauthner cell has an axon that crosses over, innervating neurons at the same brain level and then travelling down through the spinal cord, making numerous connections as it goes. The synapses generated by a Mauthner cell are so powerful that a single action potential gives rise to a major behavioral response: within milliseconds the fish curves its body into a C-shape, then straightens, thereby propelling itself rapidly forward. Functionally this is a fast escape response, triggered most easily by a strong sound wave or pressure wave impinging on the lateral line organ of the fish. Mauther cells are not the only identified neurons in fish—there are about 20 more types, including pairs of "Mauthner cell analogs" in each spinal segmental nucleus. Although a Mauthner cell is capable of bringing about an escape response all by itself, in the context of ordinary behavior other types of cells usually contribute to shaping the amplitude and direction of the response.

Mauthner cells have been described as command neurons. A command neuron is a special type of identified neuron, defined as a neuron that is capable of driving a specific behavior all by itself.[29] Such neurons appear most commonly in the fast escape systems of various species—the squid giant axon and squid giant synapse, used for pioneering experiments in neurophysiology because of their enormous size, both participate in the fast escape circuit of the squid. The concept of a command neuron has, however, become controversial, because of studies showing that some neurons that initially appeared to fit the description were really only capable of evoking a response in a limited set of circumstances

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