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Axon
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For other uses, see Axon (disambiguation).
Axon
An axon of a multipolar neuron
Identifiers
MeSH
Axons
Anatomical terminology
[edit on Wikidata]
An axon (from Greek ἄξων áxōn, axis) or nerve fiber, is a long, slender projection of a nerve cell, or neuron, that typically conducts electrical impulses known as action potentials, away from the nerve cell body. The function of the axon is to transmit information to different neurons, muscles, and glands. In certain sensory neurons (pseudounipolar neurons), such as those for touch and warmth, the axons are called afferent nerve fibers and the electrical impulse travels along these from the periphery to the cell body, and from the cell body to the spinal cord along another branch of the same axon. Axon dysfunction has caused many inherited and acquired neurological disorders which can affect both the peripheral and central neurons. Nerve fibers are classed into three types – group A nerve fibers, group B nerve fibers, and group C nerve fibers. Groups A and B are myelinated, and group C are unmyelinated. These groups include both sensory fibers and motor fibers. Another classification, groups only the sensory fibers, and these are grouped as Type I, Type II, Type III , and Type IV.
An axon is one of two types of cytoplasmic protrusions from the cell body of a neuron; the other type is a dendrite. Axons are distinguished from dendrites by several features, including shape (dendrites often taper while axons usually maintain a constant radius), length (dendrites are restricted to a small region around the cell body while axons can be much longer), and function (dendrites receive signals whereas axons transmit them). Some types of neurons have no axon and transmit signals from their dendrites. No neuron ever has more than one axon; however in invertebrates such as insects or leeches the axon sometimes consists of several regions that function more or less independently of each other.[1]
Axons are covered by a membrane known as an axolemma; the cytoplasm of an axon is called axoplasm. Most axons branch, in some cases very profusely. The end branches of an axon are called telodendria. The swollen end of a telodendron is known as the axon terminal which joins the dendron or cell body of another neuron forming a synaptic connection. Axons make contact with other cells—usually other neurons but sometimes muscle or gland cells—at junctions called synapses. In some circumstances, the axon of one neuron may form a synapse with the dendrites of the same neuron, resulting in an autapse. At a synapse, the membrane of the axon closely adjoins the membrane of the target cell, and special molecular structures serve to transmit electrical or electrochemical signals across the gap. Some synaptic junctions appear along the length of an axon as it extends—these are called en passant ("in passing") synapses and can be in the hundreds or even the thousands along one axon.[2] Other synapses appear as terminals at the ends of axonal branches.
A single axon, with all its branches taken together, can innervate multiple parts of the brain and generate thousands of synaptic terminals. A bundle of axons make a nerve tract in the central nervous system,[3] and a fascicle in the peripheral nervous system. The largest white matter tract in the brain is the corpus callosum formed of some 20 million axons.[3]
Contents
[hide]
1 Anatomy
1.1 Axonal region
1.1.1 Axon hillock
1.1.2 Initial segment
1.2 Axonal transport
1.3 Myelination
1.4 Nodes of Ranvier
1.5 Axon terminals
2 Action potentials
3 Development and growth
3.1 Development
3.1.1 Extracellular signaling
3.1.2 Intracellular signaling
3.1.3 Cytoskeletal dynamics
3.2 Growth
4 Classification
4.1 Motor
4.2 Sensory
4.3 Autonomic
5 Clinical significance
6 History
7 See also
8 References
9 External links
Anatomy[edit]
A typical myelinated axon
A dissected human brain, showing grey matter and white matter
Axons are the primary transmission lines of the nervous system, and as bundles they form nerves. Some axons can extend up to one meter or more while others extend as little as one millimeter. The longest axons in the human body are those of the sciatic nerve, which run from the base of the spinal cord to the big toe of each foot. The diameter of axons is also variable. Most individual axons are microscopic in diameter (typically about one micrometer (µm) across). The largest mammalian axons can reach a diameter of up to 20 µm. The squid giant axon, which is specialized to conduct signals very rapidly, is close to 1 millimetre in diameter, the size of a small pencil lead. The numbers of axonal telodendria (the branching structures at the end of the axon) can also differ from one nerve fiber to the next. Axons in the central nervous system (CNS) typically show multiple telodendria, with many synaptic end points. In comparison, the cerebellar granule cell axon is characterized by a single T-shaped branch node from which two parallel fibers extend. Elaborate branching allows for the simultaneous transmission of messages to a large number of target neurons within a single region of the brain.
There are two types of axons in the nervous system: myelinated and unmyelinated axons.[4] Myelin is a layer of a fatty insulating substance, which is formed by two types of glial cells Schwann cells and oligodendrocytes. In the peripheral nervous system Schwann cells form the myelin sheath of a myelinated axon. In the central nervous system oligodendrocytes form the insulating myelin. Along myelinated nerve fibers, gaps in the myelin sheath known as nodes of Ranvier occur at evenly spaced intervals. The myelination enables an especially rapid mode of electrical impulse propagation called saltatory conduction.
The myelinated axons from the cortical neurons form the bulk of the neural tissue called white matter in the brain. The myelin gives the white appearance to the tissue in contrast to the grey matter of the cerebral cortex which contains the neuronal cell bodies. A similar arrangement is seen in the cerebellum. Bundles of myelinated axons make up the nerve tracts in the CNS. Where these tracts cross the midline of the brain to connect opposite regions they are called commissures. The largest of these is the corpus callosum that connects the two cerebral hemispheres, and this has around 20 million axons.[3]
The structure of a neuron is seen to consist of two separate functional regions, or compartments – the cell body together with the dendrites as one region, and the axonal region as the other.[2] Nissl bodies of the soma and dendrites, where protein is synthesised is absent in the axonal region which includes the axon hillock.[2]
Axonal region[edit]
The axonal region or compartment, includes the axon hillock, the initial segment, the rest of the axon, and the axon telodendria, and axon terminals. It also includes the myelin sheath. The Nissl bodies that produce the neuronal proteins are absent in the axonal region.[2] Proteins needed for the growth of the axon, and the removal of waste materials, need a framework for transport. This axonal transport is provided for in the axoplasm.
Axon hillock[edit]
Detail showing microtubules at axon hillock and initial segment.
The axon hillock is the area formed from the cell body of the neuron as it extends to become the axon. It precedes the initial segment. The received action potentials that are summed in the neuron are transmitted to the axon hillock for the generation of an action potential from the initial segment.
Initial segment[edit]
The axon initial segment — the thick, unmyelinated part of an axon that connects directly to the cell body — consists of a specialized complex of proteins. It is approximately 25 µm in length and functions as the site of action potential initiation.[5] The density of voltage-gated sodium channels is much higher in the initial segment than in the remainder of the axon or in the adjacent cell body, excepting the axon hillock.[6] The voltage-gated ion channels are known to be found within certain areas of the axonal membrane and initiate action potential, conduction, and synaptic transmission.[4]
Axonal transport[edit]
The axoplasm is the equivalent of cytoplasm in the cell. Microtubules form in the axoplasm at the axon hillock. They are arranged along the length of the axon, in overlapping sections, and all point in the same direction – towards the axon terminals.[7] This is noted by the positive endings of the microtubules. This overlapping arrangement provides the routes for the transport of different materials from the cell body.[7] Studies on the axoplasm has shown the movement of numerous vesicles of all sizes to be seen along cytoskeletal filaments – the microtubules, in both directions between the axon and its terminals and the cell body.
Outgoing transport (anterograde) carries mitochondria and membrane proteins needed for growth to the axon terminal. Ingoing transport (retrograde) carries cell waste materials from the axon terminal to the soma.[8] Outgoing and ingoing tracks use different sets of motor proteins.[7] Outgoing transport is provided by kinesin, and ingoing return traffic is provided by dynein. Dinein is minus-end directed.[8] There are many forms of kinesis and dynein motor proteins, and each is thought to carry a different cargo.[7] The studies on transport in the axon led to the naming of kinesin. [7]
Myelination[edit]
Transmission electron micrograph of a myelinated axon in cross-section. Generated by the electron microscopy unit at Trinity College, Hartford CT
Cross section of an axon.
1. Axon
2. Nucleus of Schwann cell
3. Schwann cell
4. Myelin sheath
5. Neurolemma
In the nervous system, axons may be myelinated, or unmyelinated. This is the provision of an insulating layer, called a myelin sheath. In the peripheral nervous system axons are myelinated by glial cells known as Schwann cells. In the central nervous system the myelin sheath is provided by another type of glial cell, the oligodendrocyte. Schwann cells myelinate a single axon. An oligodendrocyte can myelinate up to 50 axons.[9]
Nodes of Ranvier[edit]
Main article: Node of Ranvier
Nodes of Ranvier (also known as myelin sheath gaps) are short unmyelinated segments of a myelinated axon, which are found periodically interspersed between segments of the myelin sheath. Therefore, at the point of the node of Ranvier, the axon is reduced in diameter.[10] These nodes are areas where action potentials can be generated. In saltatory conduction, electrical currents produced at each node of Ranvier are conducted with little attenuation to the next node in line, where they remain strong enough to generate another action potential. Thus in a myelinated axon, action potentials effectively "jump" from node to node, bypassing the myelinated stretches in between, resulting in a propagation speed much faster than even the fastest unmyelinated axon can sustain.
Axon terminals[edit]
An axon can divide into many branches called telodendria (Greek–end of tree). At the end of each telodendron is an axon terminal (also called a synaptic bouton, or terminal bouton). Axon terminals contain synaptic vesicles that store the neurotransmitter for release at the synapse. This makes multiple synaptic connections with other neurons possible. Sometimes the axon of a neuron may synapse onto dendrites of the same neuron, when it is known as an autapse.
Action potentials[edit]
Main article: Action potential
Structure of a typical chemical synapse
Postsynaptic
density
Voltage-
gated Ca++
channel
Synaptic
vesicle
Neurotransmitter
transporter
Receptor
Neurotransmitter
Axon terminal
Synaptic cleft
Dendrite
Most axons carry signals in the form of action potentials, which are discrete electrochemical impulses that travel rapidly along an axon, starting at the cell body and terminating at points where the axon makes synaptic contact with target cells. The defining characteristic of an action potential is that it is "all-or-nothing" — every action potential that an axon generates has essentially the same size and shape. This all-or-nothing characteristic allows action potentials to be transmitted from one end of a long axon to the other without any reduction in size. There are, however, some types of neurons with short axons that carry graded electrochemical signals, of variable amplitude.
When an action potential reaches a presynaptic terminal, it activates the synaptic transmission process. The first step is rapid opening of calcium ion channels in the membrane of the axon, allowing calcium ions to flow inward across the membrane. The resulting increase in intracellular calcium concentration causes synaptic vesicles (tiny containers enclosed by a lipid membrane) filled with a neurotransmitter chemical to fuse with the axon's membrane and empty their contents into the extracellular space. The neurotransmitter is released from the presynaptic nerve through exocytosis. The neurotransmitter chemical then diffuses across to receptors located on the membrane of the target cell. The neurotransmitter binds to these receptors and activates them. Depending on the type of receptors that are activated, the effect on the target cell can be to excite the target cell, inhibit it, or alter its metabolism in some way. This entire sequence of events often takes place in less than a thousandth of a second. Afterward, inside the presynaptic terminal, a new set of vesicles is moved into position next to the membrane, ready to be released when the next action potential arrives. The action potential is the final electrical step in the integration of synaptic messages at the scale of the neuron.[4]
A)pyramidal cell, interneuron, and short durationwaveform (Axon). (B) overlay of the three average waveforms; (C) average and standard error of peak-trough time for pyramidal cells interneurons, and putative axons.(D)Scatter plot of signal to noise ratios for individual units againstpeak-trough time for axons, pyramidal cells (PYR) and interneurons (INT).
Extracellular recordings of action potential propagation in axons has been demonstrated in freely moving animals. While extracellular somatic action potentials have been used to study cellular activity in freely moving animals such as place cells, axonal activity in both white and gray matter can also be recorded. Extracellular recordings of axon action potential propagation is distinct from somatic action potentials in three ways: 1. The signal has a shorter peak-trough duration (~150μs) than of pyramidal cells (~500μs) or interneurons (~250μs). 2. The voltage change is triphasic. 3. Activity recorded on a tetrode is seen on only one of the four recording wires. In recordings from freely moving rats, axonal signals have been isolated in white matter tracts including the alveus and the corpus callosum as well hippocampal gray matter.[11]
In fact, the generation of action potentials in vivo is sequential in nature, and these sequential spikes constitute the digital codes in the neurons. Although previous studies indicate an axonal origin of a single spike evoked by short-term pulses, physiological signals in vivo trigger the initiation of sequential spikes at the cell bodies of the neurons.[12][13]
In addition to propagating action potentials to axonal terminals, the axon is able to amplify the action potentials, which makes sure a secure propagation of sequential action potentials toward the axonal terminal. In terms of molecular mechanisms, voltage-gated sodium channels in the axons possess lower threshold and shorter refractory period in response to short-term pulses.[14]
Development and growth[edit]
Development[edit]
The development of the axon to its target, is one of the six major stages in the overall development of the nervous system.[15] Studies done on cultured hippocampal neurons suggest that neurons initially produce multiple neurites that are equivalent, yet only one of these neurites is destined to become the axon.[16] It is unclear whether axon specification precedes axon elongation or vice versa,[17] although recent evidence points to the latter. If an axon that is not fully developed is cut, the polarity can change and other neurites can potentially become the axon. This alteration of polarity only occurs when the axon is cut at least 10 μm shorter than the other neurites. After the incision is made, the longest neurite will become the future axon and all the other neurites, including the original axon, will turn into dendrites.[18] Imposing an external force on a neurite, causing it to elongate, will make it become an axon.[19] Nonetheless, axonal development is achieved through a complex interplay between extracellular signaling, intracellular signaling and cytoskeletal dynamics.
Extracellular signaling[edit]
The extracellular signals that propagate through the extracellular matrix surrounding neurons play a prominent role in axonal development.[20] These signaling molecules include proteins, neurotrophic factors, and extracellular matrix and adhesion molecules. Netrin (also known as UNC-6) a secreted protein, functions in axon formation. When the UNC-5 netrin receptor is mutated, several neurites are irregularly projected out of neurons and finally a single axon is extended anteriorly.[21][22][23][24] The neurotrophic factors – nerve growth factor (NGF), brain-derived neurotrophic factor (BDNF) and neurotrophin-3 (NTF3) are also involved in axon development and bind to Trk receptors.[25]
The ganglioside-converting enzyme plasma membrane ganglioside sialidase (PMGS), which is involved in the activation of TrkA at the tip of neutrites, is required for the elongation of axons. PMGS asymmetrically distributes to the tip of the neurite that is destined to become the future axon.[26]
Intracellular signaling[edit]
During axonal development, the activity of PI3K is increased at the tip of destined axon. Disrupting the activity of PI3K inhibits axonal development. Activation of PI3K results in the production of phosphatidylinositol (3,4,5)-trisphosphate (PtdIns) which can cause significant elongation of a neurite, converting it into an axon. As such, the overexpression of phosphatases that dephosphorylate PtdIns leads into the failure of polarization.[20]
Cytoskeletal dynamics[edit]
The neurite with the lowest actin filament content will become the axon. PGMS concentration and f-actin content are inversely correlated; when PGMS becomes enriched at the tip of a neurite, its f-actin content is substantially decreased.[26] In addition, exposure to actin-depolimerizing drugs and toxin B (which inactivates Rho-signaling) causes the formation of multiple axons. Consequently, the interruption of the actin network in a growth cone will promote its neurite to become the axon.[27]
Growth[edit]
Axon of 9 day old mouse with growth cone visible
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 a lamellipodium 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 (CAMs) 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, TAG-1 an axonal glycoprotein[28], and MAG 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.
It has also been discovered through research that if the axons of a neuron were damaged, as long as the soma (the cell body of a neuron) is not damaged, the axons would regenerate and remake the synaptic connections with neurons with the help of guidepost cells. This is also referred to as neuroregeneration.[29]
Nogo-A is a type of neurite outgrowth inhibitory component that is present in the central nervous system myelin membranes (found in an axon). It has a crucial role in restricting axonal regeneration in adult mammalian central nervous system. In recent studies, if Nogo- A is blocked and neutralized, it is possible to induce long-distance axonal regeneration which leads to enhancement of functional recovery in rats and mouse spinal cord. This has yet to be done on humans.[30] A recent study has also found that macrophages activated through a specific inflammatory pathway activated by the Dectin-1 receptor are capable of promoting axon recovery, also however causing neurotoxicity in the neuron.[31]
Classification[edit]
The axons of neurons in the human peripheral nervous system can be classified based on their physical features and signal conduction properties. Axons were known to have different thicknesses (from 0.1 to 20 µm )[2] and these differences were thought to relate to the speed that an action potential could travel along the axon – its conductance velocity. Erlanger and Gasser proved this hypothesis, and identified several types of nerve fiber, establishing a relationship between the diameter of an axon and its conductance velocity. They published their findings in 1941 giving the first classification of axons.
Axons are classified in two systems. The first one introduced by Erlanger and Gasser, grouped the fibers into three main groups using the letters A, B, and C. These groups, group A, group B, and group C include both the sensory fibers (afferents) and the motor fibres (efferents). The first group A, was subdivided into alpha, beta, gamma, and delta fibers - Aα, Aβ, Aγ, and Aδ. The motor neurons of the different motor fibers, were the lower motor neurons – alpha motor neuron, beta motor neuron, and gamma motor neuron having the Aα, Aβ, and Aγ nerve fibers respectively.
Later findings by other researchers, identified two groups of Aa fibers that were motor fibers. These were then introduced into a system that only included sensory fibers (though some of these were mixed nerves and were also motor fibers). This system refers to the sensory groups as Types and uses Roman numerals,- Type Ia, Type Ib, Type II, Type III, and Type IV.
Motor[edit]
Lower motor neurons have two kind of fibers:
Motor fiber types
Type
Erlanger-Gasser
Classification
Diameter (µm)
Myelin
Conduction velocity
Associated muscle fibers
α
Aα 13-20 Yes 80–120 m/s Extrafusal muscle fibers
β
Aβ
γ
Aγ 5-8 Yes 4–24 m/s[32][33] Intrafusal muscle fibers
Sensory[edit]
Different sensory receptors innervate different types of nerve fibers. Proprioceptors are innervated by type Ia, Ib and II sensory fibers, mechanoreceptors by type II and III sensory fibers and nociceptors and thermoreceptors by type III and IV sensory fibers.
Sensory fiber types
Type
Erlanger-Gasser
Classification
Diameter (µm)
Myelin
Conduction
velocity
Associated sensory receptors
Proprioceptors
Mechanoceptors
Nociceptors and
thermoreceptors
Ia
Aα 13-20 Yes 80–120 m/s Primary receptors of muscle spindle ✔
Ib
Aα 13-20 Yes 80–120 m/s Golgi tendon organ
II
Aβ 6-12 Yes 33–75 m/s Secondary receptors of muscle spindle
All cutaneous mechanoreceptors ✔
III
Aδ 1-5 Thin 3–30 m/s Free nerve endings of touch and pressure
Nociceptors of lateral spinothalamic tract
Cold thermoreceptors ✔
IV
C 0.2-1.5 No 0.5-2.0 m/s Nociceptors of anterior spinothalamic tract
Warmth receptors
Autonomic[edit]
The autonomic nervous system has two kinds of peripheral fibers:
Fiber types
Type
Erlanger-Gasser
Classification
Diameter (µm)
Myelin[34]
Conduction velocity
preganglionic fibers
B 1–5 Yes 3–15 m/s
postganglionic fibers
C 0.2–1.5 No 0.5–2.0 m/s
Clinical significance[edit]
Main articles: Nerve injury, Peripheral neuropathy, and Demyelinating disease
In order of degree of severity, injury to a nerve can be described as neurapraxia, axonotmesis, or neurotmesis. Concussion is considered a mild form of diffuse axonal injury.[35] The dysfunction of axons in the nervous system is one of the major causes of many inherited neurological disorders that affect both peripheral and central neurons.[4]
Demyelination of axons causes the multitude of neurological symptoms found in the disease multiple sclerosis.
History[edit]
German anatomist Otto Friedrich Karl Deiters is generally credited with the discovery of the axon by distinguishing it from the dendrites.[4] Swiss Rüdolf Albert von Kölliker and German Robert Remak were the first to identify and characterize the axon initial segment. Kölliker named the axon in 1896.[36] 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 to 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. Louis-Antoine Ranvier was the first to describe the gaps or nodes found on axons and for this contribution these axonal features are now commonly referred to as the nodes of Ranvier. Santiago Ramón y Cajal, a Spanish anatomist, proposed that axons were the output components of neurons, describing their functionality.[4] Erlanger and Gasser earlier developed the classification system for peripheral nerve fibers,[37] based on axonal conduction velocity, myelination, 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.
The longfin inshore squid has been used as a model organism – it has the longest known axon.
See also[edit]
Axon guidance
Electrophysiology
Pioneer axon
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Jump up ^ Kunik, D (2011). "Laser-based single-axon transection for high-content axon injury and regeneration studies". PLoS ONE. 6 (11): e26832. doi:10.1371/journal.pone.0026832. PMC 3206876  . PMID 22073205.
Jump up ^ Schwab, Martin E. (2004). "Nogo and axon regeneration". Current Opinion in Neurobiology. 14 (1): 118–124. doi:10.1016/j.conb.2004.01.004. PMID 15018947.
Jump up ^ Gensel; et al. (2009). "Macrophages promote axon regeneration with concurrent neurotoxicity". Journal of Neuroscience. 29 (12): 3956–3968. doi:10.1523/JNEUROSCI.3992-08.2009. PMC 2693768  . PMID 19321792.
Jump up ^ Andrew, B. L.; Part, N. J. (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 (2): 213–225. PMID 4482075.
Jump up ^ Russell, N. J. (1980). "Axonal conduction velocity changes following muscle tenotomy or deafferentation during development in the rat". J Physiol. 298: 347–360. PMC 1279120  . PMID 7359413.
Jump up ^ Pocock, Gillian; et al. (2004). Human Physiology (Second ed.). New York: Oxford University Press. pp. 187–189. ISBN 0-19-858527-6.
Jump up ^ eMedicine - Traumatic Brain Injury: Definition, Epidemiology, Pathophysiology : Article by Segun T Dawodu, MD, FAAPMR, FAANEM, CIME, DipMI(RCSed)
Jump up ^ Finger, Stanley (1994). Origins of neuroscience : a history of explorations into brain function. Oxford University Press. p. 47. ISBN 9780195146943. OCLC 27151391. Kölliker would give the "axon" its name in 1896.
Jump up ^ Sansom B, "Reflex Isolation" http://www.sansomnia.com
External links[edit]
Histology image: 3_09 at the University of Oklahoma Health Sciences Center - "Slide 3 Spinal cord"
- Bialowas, Andrzej, Carlier, Edmond, Campanac, Emilie, Debanne, Dominique, Alcaraz. Axon Physiology, GisèlePHYSIOLOGICAL REVIEWS, V. 91 (2), 04/2011, p. 555-602.
[hide]
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GND: 4200576-0
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text:
2014 Winter Olympics medal table
From Wikipedia, the free encyclopedia
Jump to: navigation, search
World map showing the medal achievements of each country during the 2014 Winter Olympics
Legend:
Gold represents countries that won at least one gold medal
Silver represents countries that won at least one silver medal
Bronze represents countries that won at least one bronze medal
Red represents countries that did not win any medals
Grey represents countries that did not participate
The 2014 Winter Olympics, officially known as the XXII Olympic Winter Games, were a winter multi-sport event held in Sochi, Russia, from 7 to 23 February. A total of 2,873 athletes from 88 nations participated in 98 events in 7 sports across 15 different disciplines.[1][2]
Initially, host nation Russia matched the Soviet Union's 1976 achievement of thirteen gold medals,[α] but 4 gold medals (13 overall) were stripped later due to doping. Norway achieved the leading position in the medal table on 24 November 2017, when Russia was stripped of two gold medals in bobsleigh.[β] However, at the end of January 2018, the Court of Arbitration for Sport cleared 28 Russian athletes and 9 out of 13 medals (including 3 gold) were reinstated, allowing Russia to return to the top position.[7]
The Netherlands achieved four podium sweeps in the speed skating, dominating the men's 500 metres, men's 5,000 metres, men's 10,000 metres, and women's 1,500 metres, surpassing the previous record of two podium sweeps.[8]
Slovenia won its first Winter Olympics gold medal ever, in alpine skiing. This was also the first Winter Olympic gold medal tie.[9] Luger Armin Zöggeler of Italy became the first athlete to achieve six Winter Olympic medals over six consecutive games,[10] all achieved at the men's singles event.[11]
Speed skater Ireen Wüst from the Netherlands achieved five medals (two gold and three silver), more than any other athlete. Korean-born Russian short track speed skater Viktor Ahn, Norwegian cross-country skier Marit Bjørgen, and Belarusian biathlete Darya Domracheva tied for the most gold medals, with three each.[12]
Contents
[hide]
1 Medal table
2 Russian team doping case
3 Changes in medal standings
4 See also
5 Footnotes
6 References
Medal table[edit]
Part of a series on
2014 Winter Olympics
Bid process (Sochi bid)
Venues
Torch relay
Mascots
Broadcasters
Opening ceremony (flag bearers)
Chronological summary
Medal table (medalists)
World and Olympic records
Controversies (Russian doping scandal • LGBT rights protests)
Closing ceremony (flag bearers)
Paralympics
IOC
ROC
SOOC
v
t
e
From left to right: Tina Maze of Slovenia (gold), Dominique Gisin of Switzerland (gold) and Lara Gut of Switzerland (bronze) atop the women's downhill alpine skiing podium in the first Winter Olympic gold medal tie.[13]
From left to right: Jan Blokhuijsen (silver), Sven Kramer (gold) and Jorrit Bergsma (bronze) with medals they earned in the men's 5,000 metres speed skating, one of the four podium sweeps by the Netherlands.[14]
See also: List of 2014 Winter Olympics medal winners
The medal table is based on information provided by the International Olympic Committee (IOC) and is consistent with IOC convention in its published medal tables. By default, the table is ordered by the number of gold medals the athletes from a nation have won, where nation is an entity represented by a National Olympic Committee (NOC). The number of silver medals is taken into consideration next and then the number of bronze medals.
In the women's downhill event in alpine skiing, two gold medals were awarded for a first place tie. No silver medal was awarded for the event.[15] In the men's super-G alpine skiing, two bronze medals were awarded for a third place tie.[16]
Host nation (Russia)
Rank
NOC
Gold
Silver
Bronze
Total
1 Russia (RUS) 11 9 9 29
2 Norway (NOR) 11 5 10 26
3 Canada (CAN) 10 10 5 25
4 United States (USA) 9 7 12 28
5 Netherlands (NED) 8 7 9 24
6 Germany (GER) 8 6 5 19
7 Switzerland (SUI) 6 3 2 11
8 Belarus (BLR) 5 0 1 6
9 Austria (AUT) 4 8 5 17
10 France (FRA) 4 4 7 15
11 Poland (POL) 4 1 1 6
12 China (CHN) 3 4 2 9
13 South Korea (KOR) 3 3 2 8
14 Sweden (SWE) 2 7 6 15
15 Czech Republic (CZE) 2 4 2 8
16 Slovenia (SLO) 2 2 4 8
17 Japan (JPN) 1 4 3 8
18 Finland (FIN) 1 3 1 5
19 Great Britain (GBR) 1 1 2 4
20 Ukraine (UKR) 1 0 1 2
21 Slovakia (SVK) 1 0 0 1
22 Italy (ITA) 0 2 6 8
23 Latvia (LAT) 0 2 2 4
24 Australia (AUS) 0 2 1 3
25 Croatia (CRO) 0 1 0 1
26 Kazakhstan (KAZ) 0 0 1 1
Total (26 NOCs)
97
95
99
291
Russian team doping case[edit]
Main articles: Doping in Russia, McLaren Report, and Oswald Commission
On 18 July 2016, the McLaren Report was published claiming that the Russian government had sanctioned the use of performance-enhancing drugs by Russian athletes in the 2014 Winter Olympics.[17]
On 9 December 2016, a World Anti-Doping Agency report expanded upon the previous report and included the note that "Two [Russian] [sport] athletes, winners of 4 Sochi Olympic Gold medals, and a female Silver medal winner in [sport] had samples with salt readings that were physiologically impossible" and that "Twelve [Russian] medal winning athletes ... from 44 examined samples had scratches and marks on the inside of the caps of their B sample bottles, indicating tampering".[18]
In December 2016, following the release of the McLaren Report on Russian doping at the Sochi Olympics, the International Olympic Committee announced the initiation of an investigation of 28 Russian athletes at the Sochi Olympic Games. La Gazzetta dello Sport reported the names of 17 athletes, of whom 15 were among the 28 under investigation (the number later rose to 46). [19][20]
Three ladies artistic skaters were named as being under investigation. They are Adelina Sotnikova, the singles gold medalist, as well as pairs skaters Tatiana Volosozhar and Ksenia Stolbova. Volosozhar and Stolbova won gold and silver medals, respectively, in pairs skating. Both also won gold medals in the team event, which also puts the other eight team medalists at risk of losing their golds.[21] In November 2017 the proceeding against Sotnikova was dropped.[22]
Six cross-country skiers were suspended from competition on the basis of the McLaren Report: Evgeniy Belov, Alexander Legkov, Alexey Petukhov, Maxim Vylegzhanin, Yulia Ivanova and Evgenia Shapovalova. Legkov won a gold and silver medals, and Vylegzhanin won three silver medals.[23] The IOC disqualified all six from Sochi, imposed lifetime bans and, in the process, stripped Legkov's and Vylegzhanin's of the medals they had won in four events (three individual medals and one team medal).[24] On 1 February 2018, the Court of Arbitration for Sport upheld five out of six appeals (except for Ivanova whose appeal was overturned), and their results were restored.[25] Nikita Kryukov, Alexander Bessmertnykh and Natalya Matveyeva were disqualified on 22 December 2017.[26] These decisions were also overturned by the CAS on 1 February 2018.[25] The IOC intends to appeal these decisions.[27]
The International Biathlon Union suspended two Russian biathletes who were in the Sochi games: Olga Vilukhina and Yana Romanova. Vilukhina won silver in sprint, and both women were on a relay team that won the silver medal.[28] They were disqualified and stripped of their medals on 27 November 2017.[29]
The International Bobsleigh and Skeleton Federation suspended four Russian skeleton sliders. They were Alexander Tretyakov, Elena Nikitina, Maria Orlova and Olga Potylitsina. Tretyakov won a gold medal, and Nikitina won a bronze.[30][31] On 22 November 2017, the IOC stripped these medals and imposed lifetime Olympic bans on all four.[32]
Seven Russian female ice hockey players were to have hearings before the Oswald Commission on 22 November 2017. The identities of the seven players have not been revealed. Two of the seven are accused of submitting samples showing readings that were physically impossible to be held by a woman. The Russian women's ice hockey team finished sixth at Sochi 2014.[33] On December 12, 2017 six of them were disqualified.[34] Tatiana Burina and Anna Shukina were also disqualified ten days later.[35]
On 24 November 2017, the IOC disqualified and imposed lifetime Olympic bans on bobsledder Alexandr Zubkov who won two gold medals and speed skater Olga Fatkulina who won a silver medal.[36] All their results were disqualified, resulting in Russia losing its first place in the medal standings. On 27 November 2017, Russian bobsledders Aleksei Negodaylo, Dmitry Trunenkov, and skeleton racer Sergei Chudinov were sanctioned as well.[37] Three other athletes who didn't win medals (Alexander Kasjanov, Ilvir Huzin, Aleksei Pushkarev) were banned on November 29, 2017.[38] Biathlete Olga Zaitseva who won silver in a relay was disqualified on 1 December 2017. Two other athletes, Anastasia Dotsenko and Yuliya Chekalyova, were also banned, bringing the total to 31 (six Russian ice hockey players were sanctioned shortly after).[39] Speed skaters Ivan Skobrev and Artyom Kuznetsov, lugers Albert Demchenko and Tatiana Ivanova, bobsledders Liudmila Udobkina and Maxim Belugin were disqualified on 22 December 2017, bringing the total to 43. Demchenko and Tatiana Ivanova were also stripped of their silver medals.[35]
As of January 2018, the IOC had sanctioned 43 Russian athletes from the 2014 Winter Olympics and banned them from competing in the 2018 edition and all other future Olympic Games as part of the Oswald Commission. All but one of these athletes appealed against their bans to the Court of Arbitration for Sport. The court overturned the sanctions on 28 athletes meaning that their Sochi medals and results are reinstated but decided that there was sufficient evidence against 11 athletes to uphold their Sochi sanctions. The IOC said in a statement that “the result of the CAS decision does not mean that athletes from the group of 28 will be invited to the Games. Not being sanctioned does not automatically confer the privilege of an invitation” and that “this [case] may have a serious impact on the future fight against doping”. The IOC found it important to note that CAS Secretary General "insisted that the CAS decision does not mean that these 28 athletes are innocent” and that they would consider an appeal against the courts decision. The court also decided that none of the 39 athletes should be banned from all future Olympic Games, but only the 2018 Games. 3 Russian athletes are still waiting for their hearing which will be conducted after the 2018 Games.[27]
Changes in medal standings[edit]
See also: List of stripped Olympic medals
Ruling date
Sport / event
NOC
01 !
02 !
03 !
Total
Comment
List of official changes in medal standings (after the Games)
1 November 2017 Cross-country skiing
Men's 50 kilometre freestyle
Men's 4 × 10 kilometre relay Russia (RUS) −1 −1 −2 On 1 November 2017, the IOC disqualified cross country skier Alexander Legkov and he was stripped of his gold and silver medals.[40]
9 November 2017 Cross-country skiing
Men's 50 kilometre freestyle
Men's team sprint Russia (RUS) −2 −2 On 9 November 2017, the IOC disqualified cross country skier Maxim Vylegzhanin and he was stripped of his two silver medals (alongside with the stripped silver medal in the team relay with Legkov).[41]
22 November 2017 Skeleton
Men's event
Women's event Russia (RUS) −1 −1 −2 On 22 November 2017, the IOC disqualified men's gold medallist Alexander Tretyakov and women's bronze medallist Elena Nikitina.[32]
24 November 2017 Bobsleigh
Two-man
Four-man Russia (RUS) –2 −2 On 24 November 2017, the IOC disqualified bobsledder Alexandr Zubkov and he was stripped of his two gold medals.[36] His teammates in four-man bobsled Alexey Negodaylo and Dmitry Trunenkov were disqualified three days later.[37] On 18 December 2017, Zubkov's teammate in two-man bobsled and four-man bobsled, Alexey Voyevoda was also disqualified.[42]
24 November 2017 Speed skating
Women's 500 metres Russia (RUS) –1 −1 On 24 November 2017, the IOC disqualified speedskater Olga Fatkulina and she was stripped of her silver medal.[36]
27 November 2017 Biathlon
Women's sprint Russia (RUS) –1 −1 On 27 November 2017, the IOC disqualified biathlete Olga Vilukhina and she was stripped of her silver medal.[37]
27 November 2017 Biathlon
Women's relay Russia (RUS) –1 −1 On 27 November 2017, the IOC disqualified biathletes Olga Vilukhina and Yana Romanova and they were stripped of their relay silver. On 1 December 2017, fellow team member Olga Zaitseva was also disqualified.[37]
22 December 2017 Luge
Men's singles
Team relay Russia (RUS) –2 −2 On 22 December 2017, the IOC disqualified lugers Albert Demchenko and Tatiana Ivanova who won a combined two silver medals. Their results were made void.[35]
1 February 2018 Cross-country skiing
Men's 50 kilometre freestyle
Men's team sprint
Men's 4 × 10 kilometre relay
Skeleton
Men's event
Women's event
Speed skating
Women's 500 metres
Luge
Men's singles
Team relay Russia (RUS) +2 +6 +1 +9 On 1 February 2018, the Court of Arbitration for Sport reinstated the results for medalists Aleksander Tretyakov and Elena Nikitina in skeleton, Alexander Legkov and Maxim Vylegzhanin in cross-country skiing, Albert Demchenko and Tatiana Ivanova in luge, and Olga Fatkulina in speed skating.[25]
List of official changes by country
NOC
Gold
Silver
Bronze
Total
Russia (RUS) −2 −2 −4
See also[edit]
2014 Winter Paralympics medal table
Footnotes[edit]
Jump up ^ Russia is legally considered to be the successor state of the Soviet Union.[3]
Jump up ^ The gold medal counts were previously topped by host nations the United States in 1932,[4] Norway in 1952,[5] and Canada in 2010.[6]
References[edit]
General
Specific
Jump up ^ "Sochi 2014". International Olympic Committee. Retrieved 25 August 2015.
Jump up ^ "2014 Sochi Winter Olympics: Schedules, Medals, Results". The Huffington Post. Retrieved 22 September 2015.
Jump up ^ Bühler, Konrad G. (2001). State Succession and Membership in International Organisations. Legal Aspects of International Organisation Series. Volume 38. Martinus Nijhoff Publishers. pp. 161–4. ISBN 9789041115539.
Jump up ^ "1932 Lake Placid Winter Games". International Olympic Committee. Archived from the original on 8 August 2008. Retrieved 25 August 2015.
Jump up ^ "1952 Oslo Winter Games". Sports Reference LLC. Archived from the original on 22 July 2011. Retrieved 25 August 2015.
Jump up ^ "Vancouver 2010 Winter Olympics–Medals". ESPN. Retrieved 25 August 2015.
Jump up ^ "28 Russians have Olympic doping bans lifted". nbcsports.com. 1 February 2018. Retrieved 4 February 2018.
Jump up ^ "Bergsma breaks Olympic record to lead fourth Dutch medal sweep". Xinhuanet. 19 February 2014. Retrieved 25 August 2015.
Jump up ^ Herman, Martyn (12 February 2014). "Maze amazes as she wins Slovenia's first gold". Reuters. Rosa Khutor, Russia. Retrieved 12 February 2014.
Jump up ^ Macur, Juliet (8 February 2014). "The Winning Formula of Luge's 'Old Man'". The New York Times. Retrieved 30 August 2015.
Jump up ^ "Armin Zöggeler". International Olympic Committee. Retrieved 22 September 2015.
Jump up ^ "2014 Sochi Winter Games". Sports Reference LLC. Retrieved 25 August 2015.
Jump up ^ "Women's downhill results". International Olympic Committee. Retrieved 23 September 2015.
Jump up ^ "Sochi 2014: Sven Kramer defends 5,000 m speed skating title". BBC Sport. 8 February 2014. Retrieved 25 August 2015.
Jump up ^ Pennington, Bill (12 February 2014). "In Women's Downhill, a Nice Round Historic Tie". The New York Times. Retrieved 12 February 2014.
Jump up ^ "Alpine skiing — Men's super-G". International Olympic Committee. Retrieved 25 August 2015.
Jump up ^ "Russia May Face Olympics Ban as Doping Scheme Is Confirmed". New York Times. 18 July 2016.
Jump up ^ "McClaren report part II" (PDF). 9 December 2016.
Jump up ^ "Ghiaccio, pattinaggio. Scandalo Sochi 2014. Sospetti sulla Sotnikova: Kostner d'argento?". La Gazzetta dello Sport. Milan, Italy. 30 December 2016. Retrieved 16 January 2017.
Jump up ^ "Media reported about the possible deprivation of the figure skater Sotnikova gold Sochi 2014". Archived from the original on 7 August 2017. Retrieved 16 January 2017.
Jump up ^ "McClaren doping report may affect Russian figure skaters". Retrieved 23 November 2017.
Jump up ^ Butler, Nick (9 November 2017). "Exclusive: Olympic figure skating champion cleared of doping charge by IOC but four Russian skiers disqualified". Retrieved 9 November 2017.
Jump up ^ "Six Russian XC Skiers and Two Biathletes Provisionally Suspended due to McLaren Report UPDATED - SkiTrax". skitrax.com. Retrieved 12 February 2018.
Jump up ^ Michael Pavitt (27 November 2017). "IOC back Rodchenkov as a reliable witness as first details of why Russians disqualified from Sochi 2014 published". insidethegames.biz. Retrieved 4 December 2017.
^ Jump up to: a b c http://www.tas-cas.org/fileadmin/user_upload/Media_Release__decision_RUS_IOC_.pdf
Jump up ^ "IOC sanctions 11 Russian athletes as part of Oswald Commission findings". International Olympic Committee. 22 December 2017. Retrieved 22 December 2017.
^ Jump up to: a b "IOC Statement on CAS Decision". International Olympic Committee. 1 February 2018. Retrieved 4 February 2018.
Jump up ^ Morgan, Liam (21 January 2017). "IBU dismisses cases against 22 Russians named in McLaren Report". Insidethegames.biz. Dunsar Media Company Ltd. Retrieved 23 November 2017.
Jump up ^ "IOC sanctions five Russian athletes and publishes first full decision as part of the Oswald Commission findings". International Olympic Committee. 2017-11-27. Retrieved 2017-11-27.
Jump up ^ "Four Russia Skeleton Athletes Suspended For Doping At 2014 Sochi Olympics". RadioFreeEurope/RadioLiberty. Retrieved 23 November 2017.
Jump up ^ "Russia's Nikitina denies knowledge of any suspension from skeleton competition". Retrieved 23 November 2017.
^ Jump up to: a b "IOC sanctions four Russian athletes as part of Oswald Commission findings". International Olympic Committee. 22 November 2017. Retrieved 22 November 2017.
Jump up ^ Morgan, Liam (21 November 2017). "FIS to provide update on Russian skiers as ice hockey players set to appear before Oswald Commission". Insidethegames.biz. Dunsar Media Company Ltd. Retrieved 22 November 2017.
Jump up ^ "IOC sanctions six Russian athletes and closes one case as part of the Oswald Commission findingsdate=December 12, 2017". olympic.org. Retrieved December 13, 2017.
^ Jump up to: a b c "IOC sanctions 11 Russian athletes as part of Oswald Commission findings". International Olympic Committee. 2017-12-22. Retrieved 2017-12-22.
^ Jump up to: a b c "IOC sanctions four Russian athletes as part of Oswald Commission findings". International Olympic Committee. 24 November 2017. Retrieved 24 November 2017.
^ Jump up to: a b c d "IOC sanctions five Russian athletes and publishes first full decision as part of the Oswald Commission findings". International Olympic Committee. 27 November 2017. Retrieved 27 November 2017.
Jump up ^ "IOC sanctions three Russian athletes as part of Oswald Commission findings". 4 December 2017. Retrieved 12 February 2018.
Jump up ^ "IOC sanctions three Russian athletes as part of Oswald Commission findings". 6 February 2018. Retrieved 12 February 2018.
Jump up ^ "IOC sanctions two Russian athletes as part of Oswald Commission findings". International Olympic Committee. 1 November 2017. Retrieved 1 November 2017.
Jump up ^ "IOC sanctions four Russian athletes and closes one case as part of Oswald Commission findings". International Olympic Committee. 9 November 2017. Retrieved 9 November 2017.
Jump up ^ "IOC sanctions one Russian athlete, and closes one case as part of Oswald Commission findings". International Olympic Committee. 19 December 2017. Retrieved 20 December 2017.
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value: Освојувачи на медали на Зимските олимписки игри 2014 – Macedonian

value:
interlanguage-link interwiki-mk
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value: Medaillespiegel van de Olympische Winterspelen 2014 – Dutch

value:
interlanguage-link interwiki-nl
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value: 2014年ソチオリンピックのメダル受賞数一覧 – Japanese

value:
interlanguage-link interwiki-ja
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value: Klasyfikacja medalowa Zimowych Igrzysk Olimpijskich 2014 – Polish

value:
interlanguage-link interwiki-pl
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value: Quadro de medalhas dos Jogos Olímpicos de Inverno de 2014 – Portuguese

value: Clasamentul pe medalii la Jocurile Olimpice de iarnă din 2014 – Romanian

value:
interlanguage-link interwiki-ro
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value: Медальный зачёт на зимних Олимпийских играх 2014 – Russian

value: 2014 Winter Olympics medal table – Scots

value:
interlanguage-link interwiki-sco
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value: Vuoden 2014 talviolympialaisten mitalitaulukko – Finnish

value:
interlanguage-link interwiki-fi
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value: Medaljfördelning vid olympiska vinterspelen 2014 – Swedish

value:
interlanguage-link interwiki-sv
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value: สรุปเหรียญโอลิมปิกฤดูหนาว 2014 – Thai

value:
interlanguage-link interwiki-th
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value: 2014 Kış Olimpiyatları madalya sıralaması – Turkish

value:
interlanguage-link interwiki-tr
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value: Таблиця медалей зимових Олімпійських ігор 2014 – Ukrainian

value:
interlanguage-link interwiki-uk
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value: تمغاجات سرمائی اولمپکس 2014ء – Urdu

value:
interlanguage-link interwiki-vep
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value: 2014年冬季奥林匹克运动会奖牌榜 – Chinese

value: All languages (initial selection from common choices by you and others)

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