Neurons

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For the journal of the same name, see Neuron (journal)

Neurons (also known as neurones, nerve cells and nerve fibers) are electrically excitable cells in the nervous system that function to process and transmit information. In vertebrate animals, neurons are the core structural components of the brain, spinal cord and peripheral nerves.

Neurons are typically composed of a soma, or cell body, a dendritic tree and an axon. The majority of vertebrate neurons receive input on the cell body and dendritic tree, and transmit output via the axon. However, there is great heterogeneity throughout the nervous system and the animal kingdom, in the size, shape and function of neurons.

Neurons communicate via chemical and electrical synapses, in a process known as synaptic transmission. The fundamental process that triggers synaptic transmission is the action potential, a propagating electrical signal that is generated by exploiting the electrically excitable membrane of the neuron.

HistoryEdit

The neuron's role as the primary functional unit of the nervous system was first recognised in the early 20th century through the work of the Spanish anatomist Santiago Ramón y Cajal. Cajal proposed that neurons were discrete cells that communicated with each other via specialized junctions, or spaces, between cells. This became known as the neuron doctrine, one of the central tenets of modern neuroscience. To observe the structure of individual neurons, Cajal used a silver staining method developed by his rival, Camillo Golgi. The Golgi stain is an extremely useful method for neuroanatomical investigations because, for reasons unknown, it stains a very small percentage of cells in a tissue, so one is able to see the complete microstructure of individual neurons without much overlap from other cells in the densely packed brain.

Anatomy and histologyEdit

Neuron
Structure of a typical neuron

Neurons are highly specialized for the processing and transmission of cellular signals. Given the diversity of functions performed by neurons in different parts of the nervous system, there is, as expected, a wide variety in the shape, size, and electrochemical properties of neurons. For instance, the soma of a neuron can vary in size from 4 to 100 micrometers in diameter. [1]

• The soma, is the central part of the neuron. It contains the nucleus of the cell, and therefore is where most protein synthesis occurs. The nucleus ranges from 3 to 18 micrometers in diameter. [2]
• The dendrites of a neuron are cellular extensions with many branches, and are referred to, therefore, as a dendritic tree. The overall shape and structure of a neuron's dendrites is called its dendritic tree, and is where the majority of input to the neuron occurs. However, information outflow (i.e. from dendrites to other neurons) can also occur.
• The axon is a finer, cable-like projection which can extend tens, hundreds, or even tens of thousands of times the diameter of the soma in length. The axon carries nerve signals away from the soma (and also carry some types of information back to it). Many neurons have only one axon, but this axon may - and usually will - undergo extensive branching, enabling communication with many target cells. The part of the axon where it emerges from the soma is called the 'axon hillock'. Besides being an anatomical structure, the axon hillock is also the part of the neuron that has the greatest density of voltage-dependent sodium channels. This makes it the most easily-excited part of the neuron and the spike initiation zone for the axon: in neurological terms it has the greatest hyperpolarized action potential threshold. While the axon and axon hillock are generally involved in information outflow, this region can also receive input from other neurons as well.
• The axon terminal is a specialized structure at the end of the axon that is used to release neurotransmitter chemicals and communicate with target neurons.

Although the canonical view of the neuron attributes dedicated functions to its various anatomical components, dendrites and axons often act in ways contrary to their so-called main function.

Axons and dendrites in the central nervous system are typically only about a micrometer thick, while some in the peripheral nervous system are much thicker. The soma is usually about 10–25 micrometers in diameter and often is not much larger than the cell nucleus it contains. The longest axon of a human motoneuron can be over a meter long, reaching from the base of the spine to the toes. Sensory neurons have axons that run from the toes to the dorsal columns, over 1.5 meters in adults. Giraffes have single axons several meters in length running along the entire length of their necks. Much of what is known about axonal function comes from studying the squid giant axon, an ideal experimental preparation because of its relatively immense size (0.5–1 millimeters thick, several centimeters long)..

ClassesEdit

Structural classificationEdit

Most neurons can be anatomically characterized as:

• Unipolar or Pseudounipolar: dendrite and axon emerging from same process.
• Bipolar: single axon and single dendrite on opposite ends of the soma.
• Multipolar: more than two dendrites
• Golgi I: neurons with long-projecting axonal processes.
• Golgi II: neurons whose axonal process projects locally.

Some unique neuronal types can be identified according to their location in the nervous system and distinct shape. Some examples are basket, Betz, medium spiny, Purkinje, pyramidal and Renshaw cells.

Functional classificationEdit

• Afferent neurons convey information from tissues and organs into the central nervous system.
• Efferent neurons transmit signals from the central nervous system to the effector cells and are sometimes called motor neurons.
• Interneurons connect neurons within specific regions of the central nervous system. These are the most abundant neurons.

Afferent and efferent can also refer generally to neurons which, respectively, bring information to or send information from brain region.

Classification by action on other neurons

• Excitatory neurons excite their target neurons. Excitatory neurons in the brain are often glutamatergic. Spinal motoneurons, which synapse on muscle cells, use acetylcholine as their neurotransmitter.
• Inhibitory neurons inhibit their target neurons. Inhibitory neurons are often interneurons. The output of some brain structures (neostriatum, globus pallidus, cerebellum) are inhibitory. The primary inhibitory neurotransmitters are GABA and glycine.
• Modulatory neurons evoke more complex effects termed neuromodulation. These neurons use such neurotransmitters as dopamine, acetylcholine, serotonin and others.

Classification by discharge patterns
Neurons can be classified according to their electrophysiological characteristics:

• Tonic or regular spiking. Some neurons are typically constantly (or tonically) active. Example: interneurons in neurostriatum.
• Phasic or bursting. Neurons that fire in bursts are called phasic.
• Fast spiking. Some neurons are notable for their fast firing rates, for example some types of cortical inhibitory interneurons, cells in globus pallidus.
• Thin-spike. Action potentials of some neurons are more narrow compared to the others. For example, interneurons in prefrontal cortex are thin-spike neurons.

Classification by neurotransmitter released

Some examples are cholinergic, GABAergic, glutamatergic and dopaminergic neurons.

ConnectivityEdit

Main article: Synapse

Neurons communicate with one another via synapses, where the axon terminal of one cell impinges upon a dendrite or soma of another (or less commonly to an axon). Neurons such as Purkinje cells in the cerebellum can have over 1000 dendritic branches, making connections with tens of thousands of other cells; other neurons, such as the magnocellular neurons of the supraoptic nucleus, have only one or two dendrites, each of which receives thousands of synapses. Synapses can be excitatory or inhibitory and will either increase or decrease activity in the target neuron. Some neurons also communicate via electrical synapses, which are direct, electrically-conductive junctions between cells.

In a chemical synapse, the process of synaptic transmission is as follows: when an action potential reaches the axon terminal, it opens voltage-gated calcium channels, allowing calcium ions to enter the terminal. Calcium causes synaptic vesicles filled with neurotransmitter molecules to fuse with the membrane, releasing their contents into the synaptic cleft. The neurotransmitters diffuse across the synaptic cleft and activate receptors on the postsynaptic neuron.

The human brain has a huge number of synapses. Each of the 1012 neurons (1,000 billion, i.e. 1 trillion) has on average 7,000 synaptic connections to other neurons. Most authors estimate that the brain of a three-year-old child has about 1016 synapses (10,000 trillion). This number declines with age, stabilizing by adulthood. Estimates vary for an adult, ranging from 1015 to 5 x 1015 synapses (1,000 to 5,000 trillion). [3]

The cell membrane in the axon and soma contain voltage-gated ion channels which allow the neuron to generate and propagate an electrical impulse (an action potential). Substantial early knowledge of neuron electrical activity came from experiments with squid giant axons. In 1937, John Zachary Young suggested that the giant squid axon can be used to study neuronal electrical properties. [4] As they are much larger than human neurons, but similar in nature, it was easier to study them with the technology of that time. By inserting electrodes into the giant squid axons, accurate measurements could be made of the membrane potential.

Electrical activity can be produced in neurons by a number of stimuli. Pressure, stretch, chemical transmitters, and electrical current passing across the nerve membrane as a result of a difference in voltage can all initiate nerve activity. [5]

The narrow cross-section of axons lessens the metabolic expense of carrying action potentials, but thicker axons convey impulses more rapidly. To minimize metabolic expense while maintaining rapid conduction, many neurons have insulating sheaths of myelin around their axons. The sheaths are formed by glial cells: oligodendrocytes in the central nervous system and Schwann cells in the peripheral nervous system. The sheath enables action potentials to travel faster than in unmyelinated axons of the same diameter, whilst using less energy. The myelin sheath in peripheral nerves normally runs along the axon in sections about 1 mm long, punctuated by unsheathed nodes of Ranvier which contain a high density of voltage-gated ion channels. Multiple sclerosis is a neurological disorder that results from abnormal demyelination of peripheral nerves. Neurons with demyelinated axons do not conduct electrical signals properly.

Some neurons do not generate action potentials, but instead generate a graded electrical signal, which in turn causes graded neurotransmitter release. Such nonspiking neurons tend to be sensory neurons or interneurons, because they cannot carry signals long distances.

Histology and internal structure Edit

Nerve cell bodies stained with basophilic dyes show numerous microscopic clumps of Nissl substance (named after German psychiatrist and neuropathologist Franz Nissl, 1860–1919), which consists of rough endoplasmic reticulum and associated ribosomes. The prominence of the Nissl substance can be explained by the fact that nerve cells are metabolically very active, and hence are involved in large amounts of protein synthesis.

The cell body of a neuron is supported by a complex meshwork of structural proteins called neurofilaments, which are assembled into larger neurofibrils. Some neurons also contain pigment granules, such as neuromelanin (a brownish-black pigment, byproduct of synthesis of catecholamines) and lipofuscin (yellowish-brown pigment that accumulates with age).

There are different internal structural characteristics between axons and dendrites. Axons typically almost never contain ribosomes, except some in the initial segment. Dendrites contain granular endoplasmic reticulum or ribosomes, with diminishing amounts with distance from the cell body.

Challenges to the neuron doctrineEdit

The neuron doctrine is a central tenet of modern neuroscience, but recent studies suggest that this doctrine needs to be revised.

First, electrical synapses are more common in the central nervous system than previously thought. Thus, rather than functioning as individual units, in some parts of the brain large ensembles of neurons may be active simultaneously to process neural information.[6]

Second, dendrites, like axons, also have voltage-gated ion channels and can generate electrical potentials that carry information to and from the soma. This challenges the view that dendrites are simply passive recipients of information and axons the sole transmitters. It also suggests that the neuron is not simply active as a single element, but that complex computations can occur within a single neuron.[7]

Third, the role of glia in processing neural information has begun to be appreciated. Neurons and glia make up the two chief cell types of the central nervous system. There are far more glial cells than neurons: glia outnumber neurons by as many as 10:1. Recent experimental results have suggested that glia play a vital role in information processing.[8]

Finally, recent research has challenged the historical view that neurogenesis, or the generation of new neurons, does not occur in adult mammalian brains. It is now known that the adult brain continuously creates new neurons in the hippocampus and in an area contributing to the olfactory bulb. This research has shown that neurogenesis is environment-dependent (eg. exercise, diet, interactive surroundings), age-related, upregulated by a number of growth factors, and halted by survival-type stress factors.[9][10] Of particularly compelling interest, Charles Gross and Elizabeth Gould provided evidence suggestive that neurogenesis occurred in neocortex after birth, in areas of the brain known to be important for cognitive function.[11] Strong challenges to this work have come from more well-controlled studies by Pasko Rakic and others which support Rakic's original hypothesis that neurogenesis after birth is restricted to the olfactory bulb and hippocampus.[12][13][14] Rakic argues that the Princeton group's work has not been substantiated by multiple other groups.[15]

Neurons in the brainEdit

The number of neurons in the brain varies dramatically from species to species. The human brain has about 100 billion ($10^{11}$) neurons and 100 trillion ($10^{14}$) synapses. By contrast, the nematode worm (Caenorhabditis elegans) has just 302 neurons making it an ideal experimental subject as scientists have been able to map all of the organism's neurons. Many properties of neurons, from the type of neurotransmitters used to ion channel composition, are maintained across species, allowing scientists to study processes occurring in more complex organisms in much simpler experimental systems.

ReferencesEdit

1. The Neuron: Size Comparison
2. Drachman D (2005). Do we have brain to spare?. Neurology 64 (12): 2004-5. PMID 15985565.
3. Milestones in Neuroscience Research
4. Electrical activity of nerves
5. Connors B, Long M. Electrical synapses in the mammalian brain.. Annu Rev Neurosci 27: 393-418. PMID 15217338.
6. Djurisic M, Antic S, Chen W, Zecevic D (2004). Voltage imaging from dendrites of mitral cells: EPSP attenuation and spike trigger zones.. J Neurosci 24 (30): 6703-14. PMID 15282273.
7. Witcher M, Kirov S, Harris K (2007). Plasticity of perisynaptic astroglia during synaptogenesis in the mature rat hippocampus.. Glia 55 (1): 13-23. PMID 17001633.
8. The reinvention of the self
9. Scientists Discover Addition of New Brain Cells in Highest Brain Area
10. Gould E, Reeves A, Graziano M, Gross C (1999). Neurogenesis in the neocortex of adult primates.. Science 286 (5439): 548-52. PMID 10521353.
11. Bhardwaj R, Curtis M, Spalding K, Buchholz B, Fink D, Björk-Eriksson T, Nordborg C, Gage F, Druid H, Eriksson P, Frisén J (2006). Neocortical neurogenesis in humans is restricted to development.. Proc Natl Acad Sci U S A 103 (33): 12564-8. PMID 16901981.
12. Rakic P (1974). Neurons in rhesus monkey visual cortex: systematic relation between time of origin and eventual disposition.. Science 183 (123): 425-7. PMID 4203022.
13. Kornack D, Rakic P (2001). Cell proliferation without neurogenesis in adult primate neocortex.. Science 294 (5549): 2127-30. PMID 11739948.
14. Rakic P (2006). Neuroscience. No more cortical neurons for you.. Science 313 (5789): 928-9. PMID 16917050.

SourcesEdit

• Kandel E.R., Schwartz, J.H., Jessell, T.M. 2000. Principles of Neural Science, 4th ed., McGraw-Hill, New York.
• Bullock, T.H., Bennett, M.V.L., Johnston, D., Josephson, R., Marder, E., Fields R.D. 2005. The Neuron Doctrine, Redux, Science, V.310, p. 791-793.
• Ramón y Cajal, S. 1933 Histology, 10th ed., Wood, Baltimore.
• Roberts A., Bush B.M.H. 1981. Neurones Without Impulses. Cambridge University Press, Cambridge.
• Peters, A., Palay, S.L., Webster, H, D., 1991 The Fine Structure of the Nervous System, 3rd ed., Oxford, New York.