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|Structure of a typical neuron|
Axons are in effect the primary transmission lines of the nervous system, and as bundles they help make up nerves. Individual axons are microscopic in diameter - typically about one micrometre across (1μm) - but may extend to macroscopic (>1mm) lengths. The longest axons in the human body, for example, are those of the sciatic nerve, which run from the base of the spine to the big toe of each foot. These single-cell fibers of the sciatic nerve may extend a meter or even longer.
In vertebrates, the axons of many neurons are sheathed in myelin, which is formed by either of two types of glial cells: Schwann cells ensheathing peripheral neurons and oligodendrocytes insulating those of the central nervous system. Along myelinated nerve fibers, gaps in the sheath known as nodes of Ranvier occur at evenly-spaced intervals, enabling an especially rapid mode of electrical impulse propagation called saltation. The demyelination of axons is what causes the multitude of neurological symptoms found in the disease Multiple Sclerosis. The axons of some neurons branch to form axon collaterals, that can be divided into a number of smaller branches called telodendria. Along these the bifurcated impulse travels simultaneously to signal more than one other cell.
Peripheral nerve fibers can be classified based on axonal conduction velocity, mylenation, fiber size etc. For example, there are slow-conducting unmyelinated C fibers and faster-conducting myelinated Aδ fibers. More complex mathematical modeling continues to be done today.
There are several types of sensory- as well as motorfibers. Other fibers not mentioned in table are e.g. fibers of the autonomic nervous system
Lower motor neurons have two kind of fibers:
|Type||Diameter||Conduction velocity||Associated muscle fibers|
|α||Extrafusal muscle fibers|
|γ||4-24 m/s||Intrafusal muscle fibers|
Different sensory receptors are innervated by different types of nerve fibers. Muscles and associated sensory receptors are innvervated by type I and II sensory fibers, while cutaneous receptors are innervated by Aβ, Aδ and C fibers.
|Type||Diameter||Conduction velocity||Associated sensory receptors|
|Ia & II||Receptors of muscle spindle|
|Ib||Golgi tendon organ|
|Aβ||6-12 µm diameter||33-75 m/s||All cutaneous mechanoreceptors|
|Aδ||1-5 µm||3-30 m/s||Free nerve endings of touch and pressure |
Nociceptors of neospinothalamic tract
|C||0.2-1.5 µm||0.5-2.0 m/s||Nociceptors of paleospinothalamic tract |
Growth and development
Growing axons move through their environment via the growth cone, which is at the tip of the axon. The growth cone has a broad sheet like extension called lamellipodia which contain protrusions called filopodia. The filopodia are the mechanism by which the entire process adheres to surfaces and explores the surrounding environment. Actin plays a major role in the mobility of this system. Environments with high levels of cell adhesion molecules or CAM's create an ideal environment for axonal growth. This seems to provide a "sticky" surface for axons to grow along. Examples of CAM's specific to neural systems include N-CAM, neuroglial CAM or NgCAM, TAG-1, MAG, and DCC, all of which are part of the immunoglobulin superfamily. Another set of molecules called extracellular matrix adhesion molecules also provide a sticky substrate for axons to grow along. Examples of these molecules include laminin, fibronectin, tenascin, and perlecan. Some of these are surface bound to cells and thus act as short range attractants or repellents. Others are difusible ligands and thus can have long range effects.
Cells called guidepost cells assist in the guidance of neuronal axon growth. These cells are typically other, sometimes immature, neurons.
Some of the first intracellular recordings in a nervous system were made in the late 1930's by K. Cole and H. Curtis. Alan Hodgkin and Andrew Huxley also employed the squid giant axon (1939) and by 1952 they had obtained a full quantitative description of the ionic basis of the action potential, leading the formulation of the Hodgkin-Huxley Model. Hodgkin and Huxley were awarded jointly the Nobel Prize for this work in 1963. The formulas detailing axonal conductance were extended to vertebrates in the Frankenhaeuser-Huxley equations. Erlanger and Gasser later developed the classification system for peripheral nerve fibers, based on axonal conduction velocity, mylenation, fiber size etc. Even recently our understanding of the biochemical basis for action potential propagation has advanced, and now includes many details about individual ion channels.
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- Andrew BL, Part NJ (1972) Properties of fast and slow motor units in hind limb and tail muscles of the rat. Q J Exp Physiol Cogn Med Sci 57:213-225.
- Russell NJ (1980) Axonal conduction velocity changes following muscle tenotomy or deafferentation during development in the rat. J Physiol 298:347-360.