text:
Different Drum
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This article is about the song written by Michael Nesmith. For the Linda Ronstadt album, see Different Drum (album). For the record label, see A Different Drum.
"Different Drum"
Single by The Stone Poneys featuring Linda Ronstadt
from the album Evergreen, Volume 2
B-side
"I've Got to Know"
Released
September 1967
Format
7" single
Genre
Baroque rock[1]
Length
2:45
Label
Capitol
Songwriter(s)
Mike Nesmith
Producer(s)
Nick Venet
The Stone Poneys featuring Linda Ronstadt singles chronology
"One for One"
(1967) "Different Drum"
(1967) "Up to My Neck in High Muddy Water"
(1968)
"One for One"
(1967) "Different Drum"
(1967) "Up to My Neck in High Muddy Water"
(1968)
"Different Drum" is a song written by Mike Nesmith in 1965 and originally recorded by the northern bluegrass band the Greenbriar Boys and included on their 1966 album, Better Late than Never!.
The song first became popular in 1967 when recorded by the Stone Poneys featuring Linda Ronstadt, who took their version of "Different Drum" to #12 on the Cash Box Top 100, #13 on the Billboard Hot 100 chart, and #16 in Record World magazine. The song did best in New Zealand, where it reached #5.[2] In 1972, Nesmith recorded his own version of his song. "Different Drum" has since been covered by other artists over the years.
Contents
[hide]
1 Mike Nesmith version
2 The Stone Poneys version
3 Chart history
3.1 Weekly charts
3.2 Year-end charts
4 Other versions
5 References
6 External links
Mike Nesmith version[edit]
The song reached a wider audience when Mike Nesmith rushed through a version of it in a comedy bit while pretending to be Billy Roy Hodstetter, in the Monkees television show episode "Too Many Girls", which aired in December 1966. Davy Jones mentions this during the commentary track on some DVDs of this episode.[3]
The song tells of a pair of lovers, one of who wants to settle down, while the other wants to retain a sense of freedom and independence. Its narrator is the lover who wants to remain free, telling the other that "we'll both live a lot longer" if they part ways now.
Nesmith later rerecorded the song for his 1972 LP And the Hits Just Keep on Comin'. His version contains four verses, as opposed to the three in Ronstadt's version.
The Stone Poneys version[edit]
The song is best known for the 1967 version credited to the Stone Poneys[4] featuring a vocal performance by a young and up-and-coming singer named Linda Ronstadt. The song was Ronstadt's first hit single, reaching number 13 on the Billboard Hot 100 as well as number 12 on the Cash Box magazine singles chart. (It went to number 1 in the Los Angeles market and number 6 in Detroit.) [5] Ronstadt's version flips the gender references in Nesmith's original lyrics, replacing "girl" with "boy" when describing her lover, but still referring to him being "pretty". The Stone Poneys had initially intended to record an "acoustic ballad version" of the song, but producer Nick Venet opted for a more complex instrumental approach, using an arrangement by Jimmy Bond (who also played bass), guitarist Al Viola, drummer Jim Gordon, strings led by Sid Sharp, and harpsichord played in baroque style (and largely improvised during the recording) by Don Randi. As a result, Ronstadt was the only member of the Stone Poneys who actually performed on the record. The album rendition offers a different stereo mix than the hit single, including a longer harpsichord bridge. [6] Ronstadt later commented that she had been surprised and "completely confused" by the changed approach to the song, and that even years later she perceived "fear and a lack of confidence" in her performance. Nesmith, on the other hand, said that Ronstadt's performance of his song "infused it with a new level of passion and sensuality".[6] In later live performances of the song, Nesmith would often sing the closing verse in the same singing style as the Ronstadt version.
Chart history[edit]
Weekly charts[edit]
Chart (1967-68)
Peak
position
Australia (Go-Set)[7] 9
Canada RPM Top Singles[8] 18
New Zealand (Listener)[9] 5
US Billboard Hot 100[10] 13
US Cash Box Top 100[11] 12
Year-end charts[edit]
Chart (1968)
Rank
Australia [12] 39
US (Joel Whitburn's Pop Annual)[13] 110
Other versions[edit]
The song has been covered by many artists.
The Poppy Family with Susan Jacks
Pete Burns
Flying Emus
Susanna Hoffs with Matthew Sweet
The Jayhawks
The Lemonheads
Me First and the Gimme Gimmes
The Pastels
Victoria Shaw
Sara Watkins
Paul Westerberg
P.P. Arnold
Gina Jeffreys
La Sera
The Lennon Sisters
Skeeter Davis
Carrie Underwood
Sarah White
References[edit]
Jump up ^ Gendron, Bernard (2002). Between Montmartre and the Mudd Club: Popular Music and the Avant-Garde. University of Chicago Press. ISBN 978-0-226-28737-9.
Jump up ^ Flavour of New Zealand, 15 March 1968
Jump up ^ "Too Many Girls". The Monkees. Season 1. Episode 15. 1966-12-19. NBC.
Jump up ^ Show 44 - Revolt of the Fat Angel: Some samples of the Los Angeles sound. [Part 4] : UNT Digital Library
Jump up ^ CMT.com : Linda Ronstadt : Biography
^ Jump up to: a b Marc Myers, "Linda Ronstadt's 'Different Drum': She and songwriter Michael Nesmith talk about her first hit", The Wall Street Journal, October 31, 2013.
Jump up ^ Go-Set National Top 40, 21 February 1968
Jump up ^ "Item Display - RPM - Library and Archives Canada". Collectionscanada.gc.ca. Retrieved 2017-05-15.
Jump up ^ Flavour of New Zealand, 15 March 1968
Jump up ^ Joel Whitburn's Top Pop Singles 1955-1990 - ISBN 0-89820-089-X
Jump up ^ Cash Box Top 100 Singles, February 3, 1968
Jump up ^ "Go-Set Magazine Charts". www.poparchives.com.au. Barry McKay. January 2007. Retrieved 10 July 2017.
Jump up ^ Whitburn, Joel (1999). Pop Annual. Menomonee Falls, Wisconsin: Record Research Inc. ISBN 0-89820-142-X.
External links[edit]
Linda Ronstadt and The Stone Poneys Alternate take on Different Drum
[show]
v
t
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Linda Ronstadt
Stone Poneys albums
The Stone Poneys featuring Linda Ronstadt
Evergreen, Volume 2
Linda Ronstadt, Stone Poneys and Friends, Vol. III
Studio albums
Hand Sown ... Home Grown
Silk Purse
Linda Ronstadt
Don't Cry Now
Heart Like a Wheel
Prisoner in Disguise
Hasten Down the Wind
Simple Dreams
Living in the USA
Mad Love
Get Closer
What's New
Lush Life
For Sentimental Reasons
Canciones de Mi Padre
Cry Like a Rainstorm, Howl Like the Wind
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Winter Light
Feels Like Home
Dedicated to the One I Love
We Ran
A Merry Little Christmas
Hummin' to Myself
Collaborations
Trio
Santiago
Trio II
Western Wall: The Tucson Sessions
Adieu False Heart
Compilations
Different Drum
Greatest Hits
Greatest Hits, Volume 2
The Very Best of Linda Ronstadt
Mi Jardin Azul: Las Canciones Favoritas
Duets
Hit singles
"Different Drum"
"Long, Long Time"
"You're No Good"
"When Will I Be Loved"
"Heat Wave"
"Love Is a Rose"
"The Tracks of My Tears"
"That'll Be the Day"
"Blue Bayou"
"It's So Easy"
"Poor Poor Pitiful Me"
"Tumbling Dice"
"Back in the U.S.A."
"Ooh Baby Baby"
"Just One Look"
"Alison"
"How Do I Make You"
"Hurt So Bad"
"I Can't Let Go"
"I Knew You When"
"What's New?"
"Somewhere Out There
"Don't Know Much"
"All My Life"
"When Something Is Wrong With My Baby"
"Walk On"
Related topics
Discography
Mr. Plow
Book
[show]
v
t
e
The Lemonheads
Evan Dando
Vess Ruhtenberg
Devon Ashley
Ben Deily
Corey Loog Brennan
John Strohm
Doug Trachten
Jesse Peretz
Juliana Hatfield
Nic Dalton
Dave Ryan
Patrick Murphy
Karl Alvarez
Bill Stevenson
P. David Hazel
Studio albums
Hate Your Friends (1987)
Creator (1988)
Lick (1989)
Lovey (1990)
It's a Shame About Ray (1992)
Come on Feel the Lemonheads (1993)
Car Button Cloth (1996)
The Lemonheads (2006)
Varshons (2009)
EPs and compilations
Favourite Spanish Dishes (1990)
The Best of the Lemonheads: The Atlantic Years (1998)
Singles
"Luka"
"Different Drum"
"Gonna Get Along Without Ya Now"
"Mrs. Robinson/Being Around"
"Into Your Arms"
Related articles
Discography
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Axon
From Wikipedia, the free encyclopedia
Jump to: navigation, search
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
References[edit]
Jump up ^ Yau, KW (1976). "Receptive fields, geometry and conduction block of sensory neurons in the CNS of the leech". J. Physiol. Lond. 263 (3): 513–538. doi:10.1113/jphysiol.1976.sp011643. PMC 1307715  . PMID 1018277.
^ Jump up to: a b c d e Squire, Larry (2013). Fundamental neuroscience (4th ed. ed.). Amsterdam: Elsevier/Academic Press. pp. 61–65. ISBN 978-0-12-385-870-2. CS1 maint: Extra text (link)
^ Jump up to: a b c Luders, E.; Thompson, P. M.; Toga, A. W. (18 August 2010). "The Development of the Corpus Callosum in the Healthy Human Brain". Journal of Neuroscience. 30 (33): 10985–10990. doi:10.1523/JNEUROSCI.5122-09.2010.
^ Jump up to: a b c d e f Debanne, Dominique; Campanac, Emilie; Bialowas, Andrzej; Carlier, Edmond; Alcaraz, Gisèle (Apr 2011). "Axon physiology". Physiological Reviews. 91 (2): 555–602. doi:10.1152/physrev.00048.2009. PMID 21527732.
Jump up ^ Clark, BD; Goldberg, EM; Rudy, B (Dec 2009). "Electrogenic tuning of the axon initial segment". The Neuroscientist : a review journal bringing neurobiology, neurology and psychiatry. 15 (6): 651–68. doi:10.1177/1073858409341973. PMC 2951114  . PMID 20007821.
Jump up ^ Wollner, DA; Catterall, WA (Nov 1986). "Localization of sodium channels in axon hillocks and initial segments of retinal ganglion cells". Proceedings of the National Academy of Sciences of the United States of America. 83 (21): 8424–8. doi:10.1073/pnas.83.21.8424. PMC 386941  . PMID 2430289.
^ Jump up to: a b c d e Alberts, Bruce (2004). Essential cell biology : an introduction to the molecular biology of the cell (2nd. ed.). New York [u.a.]: Garland. pp. 584–587. ISBN 0-8153-3481-8.
^ Jump up to: a b Alberts, Bruce (2002). Molecular biology of the cell (4th. ed.). New York [u.a.]: Garland. pp. 979–981. ISBN 0-8153-4072-9.
Jump up ^ Sadler, T. (2010). Langman's medical embryology (11th ed.). Philadelphia: Lippincott William & Wilkins. p. 300. ISBN 978-0-7817-9069-7.
Jump up ^ Hess, A; Young, JZ (November 20, 1952). "The nodes of Ranvier". Proceedings of the Royal Society. Series B. 140 (900): 301–320. doi:10.1098/rspb.1952.0063. JSTOR 82721. PMID 13003931.
Jump up ^ Robbins, A; Fox, S (Nov 2013). "Short Duration Waveforms Recorded Extracellularly from Freely Moving Rats are Representative of Axonal Activity". Frontiers in Neural Circuits. 7 (181): 7–11. doi:10.3389/fncir.2013.00181. PMC 3831546  . PMID 24348338.
Jump up ^ Rongjing Ge, Hao Qian and Jin-Hui Wang* (2011) Molecular Brain 4(19), 1~11
Jump up ^ Rongjing Ge, Hao Qian, Na Chen and Jin-Hui Wang* (2014) Molecular Brain 7(26):1-16
Jump up ^ Chen, Na; Yu, Jiandong; Qian, Hao; Jin-Hui (2010). "Axons Amplify Somatic Incomplete Spikes into Uniform Amplitudes in Mouse Cortical Pyramidal Neurons". PLOS ONE. 5 (7): e11868. doi:10.1371/journal.pone.0011868  . PMC 2912328  . PMID 20686619.
Jump up ^ Wolpert, Lewis (2015). Principles of development (5th ed.). pp. 520–524. ISBN 978-0-19-967814-3.
Jump up ^ Fletcher, TL; Banker, GA (Dec 1989). "The establishment of polarity by hippocampal neurons: the relationship between the stage of a cell's development in situ and its subsequent development in culture". Developmental Biology. 136 (2): 446–54. doi:10.1016/0012-1606(89)90269-8. PMID 2583372.
Jump up ^ Jiang, H; Rao, Y (May 2005). "Axon formation: fate versus growth". Nature Neuroscience. 8 (5): 544–6. doi:10.1038/nn0505-544. PMID 15856056.
Jump up ^ Goslin, K; Banker, G (Apr 1989). "Experimental observations on the development of polarity by hippocampal neurons in culture". The Journal of Cell Biology. 108 (4): 1507–16. doi:10.1083/jcb.108.4.1507. PMC 2115496  . PMID 2925793.
Jump up ^ Lamoureux, P; Ruthel, G; Buxbaum, RE; Heidemann, SR (Nov 11, 2002). "Mechanical tension can specify axonal fate in hippocampal neurons". The Journal of Cell Biology. 159 (3): 499–508. doi:10.1083/jcb.200207174. PMC 2173080  . PMID 12417580.
^ Jump up to: a b Arimura, N; Kaibuchi, K (Mar 2007). "Neuronal polarity: from extracellular signals to intracellular mechanisms". Nature Reviews. Neuroscience. 8 (3): 194–205. doi:10.1038/nrn2056. PMID 17311006.
Jump up ^ Neuroglia and pioneer neurons express UNC-6 to provide global and local netrin cues for guiding migrations in C. elegans
Jump up ^ Serafini, T; Kennedy, TE; Galko, MJ; Mirzayan, C; Jessell, TM; Tessier-Lavigne, M (Aug 12, 1994). "The netrins define a family of axon outgrowth-promoting proteins homologous to C. elegans UNC-6". Cell. 78 (3): 409–24. doi:10.1016/0092-8674(94)90420-0. PMID 8062384.
Jump up ^ Hong, K; Hinck, L; Nishiyama, M; Poo, MM; Tessier-Lavigne, M; Stein, E (Jun 25, 1999). "A ligand-gated association between cytoplasmic domains of UNC5 and DCC family receptors converts netrin-induced growth cone attraction to repulsion". Cell. 97 (7): 927–41. doi:10.1016/S0092-8674(00)80804-1. PMID 10399920.
Jump up ^ Hedgecock, EM; Culotti, JG; Hall, DH (Jan 1990). "The unc-5, unc-6, and unc-40 genes guide circumferential migrations of pioneer axons and mesodermal cells on the epidermis in C. elegans". Neuron. 4 (1): 61–85. doi:10.1016/0896-6273(90)90444-K. PMID 2310575.
Jump up ^ Huang, EJ; Reichardt, LF (2003). "Trk receptors: roles in neuronal signal transduction". Annual Review of Biochemistry. 72: 609–42. doi:10.1146/annurev.biochem.72.121801.161629. PMID 12676795.
^ Jump up to: a b Da Silva, JS; Hasegawa, T; Miyagi, T; Dotti, CG; Abad-Rodriguez, J (May 2005). "Asymmetric membrane ganglioside sialidase activity specifies axonal fate". Nature Neuroscience. 8 (5): 606–15. doi:10.1038/nn1442. PMID 15834419.
Jump up ^ Bradke, F; Dotti, CG (Mar 19, 1999). "The role of local actin instability in axon formation". Science. 283 (5409): 1931–4. doi:10.1126/science.283.5409.1931. PMID 10082468.
Jump up ^ Furley, AJ; Morton, SB; Manalo, D; Karagogeos, D; Dodd, J; Jessell, TM (6 April 1990). "The axonal glycoprotein TAG-1 is an immunoglobulin superfamily member with neurite outgrowth-promoting activity". Cell. 61 (1): 157–70. PMID 2317872.
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]
v
t
e
Nervous tissue
CNS
Tissue Types
Grey matter
White matter
Projection fibers
Association fiber
Commissural fiber
Lemniscus
Funiculus
Fasciculus
Nerve tract
Decussation
Commissure
Neuropil
Meninges
Cell Types
Neuronal
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insulating:
Myelination: Oligodendrocyte
other
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Postganglionic nerve fibers)
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Neurons/
nerve fibers
Parts
Soma
Axon hillock
Axon
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Types
Bipolar
Unipolar
Pseudounipolar
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Interneuron
Renshaw
Afferent nerve fiber/
Sensory neuron
GSA
GVA
SSA
SVA
fibers
Ia or Aα
Ib or Golgi or Aα
II or Aβ and Aγ
III or Aδ or fast pain
IV or C or slow pain
Efferent nerve fiber/
Motor neuron
GSE
GVE
SVE
Upper motor neuron
Lower motor neuron
α motorneuron
β motorneuron
γ motorneuron
Termination
Synapse
Electrical synapse/Gap junction
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Synaptic vesicle
Active zone
Postsynaptic density
Autapse
Ribbon synapse
Neuromuscular junction
Sensory receptors
Meissner's corpuscle
Merkel nerve ending
Pacinian corpuscle
Ruffini ending
Muscle spindle
Free nerve ending
Nociceptor
Olfactory receptor neuron
Photoreceptor cell
Hair cell
Taste bud
Authority control
GND: 4200576-0
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Standing Rules of the United States Senate
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The Standing Rules of the Senate are the parliamentary procedures adopted by the United States Senate that govern its procedure. The Senate's power to establish rules derives from Article One, Section 5 of the United States Constitution: "Each House may determine the rules of its proceedings..."
There are currently 44 rules, with the latest revision having been adopted on January 24, 2013.[1] (The Legislative Transparency and Accountability Act of 2006 lobbying reform bill introduces a 44th rule on earmarks). The stricter rules are often waived by unanimous consent.
The Constitution provides that a majority of the Senate constitutes a quorum to do business. Under the rules and customs of the Senate, a quorum is always assumed to be present unless a quorum call explicitly demonstrates otherwise. Any senator may request a quorum call by "suggesting the absence of a quorum"; a clerk then calls the roll of the Senate and notes which members are present. In practice, senators almost always request quorum calls not to establish the presence of a quorum, but to temporarily delay proceedings without having to adjourn the session. Such a delay may serve one of many purposes; often, it allows Senate leaders to negotiate compromises off the floor or to allow Senators time to come to the Senate floor to make speeches without having to constantly be present in the chamber while waiting for the opportunity. Once the need for a delay has ended, any senator may request unanimous consent to rescind the quorum call.
During debates, senators may only speak if called upon by the presiding officer. The presiding officer is, however, required to recognize the first senator who rises to speak. Thus, the presiding officer has little control over the course of debate. Customarily, the Majority Leader and Minority Leader are accorded priority during debates, even if another senator rises first. All speeches must be addressed to the presiding officer, using the words "Mr. President" or "Madam President." Only the presiding officer may be directly addressed in speeches; other Members must be referred to in the third person. In most cases, senators do not refer to each other by name, but by state, using forms such as "the senior senator from Virginia" or "the junior senator from California."
There are very few restrictions on the content of speeches; there is no requirement that speeches be germane to the matter before the Senate.
The Standing Rules of the United States Senate provide that no senator may make more than two speeches on a motion or bill on the same legislative day. (A legislative day begins when the Senate convenes and ends with adjournment; hence, it does not necessarily coincide with the calendar day.) The length of these speeches is not limited by the rules; thus, in most cases, senators may speak for as long as they please. Often, the Senate adopts unanimous consent agreements imposing time limits. In other cases (for example, for the Budget process), limits are imposed by statute. In general, however, the right to unlimited debate is preserved.
The filibuster is an obstructionary tactic used to defeat bills and motions by prolonging debate indefinitely. A filibuster may entail, but does not actually require, long speeches, dilatory motions, and an extensive series of proposed amendments. The longest filibuster speech in the history of the Senate was delivered by Strom Thurmond, who spoke for over twenty-four hours in an unsuccessful attempt to block the passage of the Civil Rights Act of 1957. The Senate may end a filibuster by invoking cloture. In most cases, cloture requires the support of three-fifths of the Senate; however, if the matter before the Senate involves changing the rules of the body, a two-thirds majority is required. Cloture is invoked very rarely, particularly because bipartisan support is usually necessary to obtain the required supermajority. If the Senate does invoke cloture, debate does not end immediately; instead, further debate is limited to thirty additional hours unless increased by another three-fifths vote.
When debate concludes, the motion in question is put to a vote. In many cases, the Senate votes by voice vote; the presiding officer puts the question, and Members respond either "Aye!" (in favor of the motion) or "No!" (against the motion). The presiding officer then announces the result of the voice vote. Any senator, however, may challenge the presiding officer's assessment and request a recorded vote. The request may be granted only if it is seconded by one-fifth of the senators present. In practice, however, senators second requests for recorded votes as a matter of courtesy. When a recorded vote is held, the clerk calls the roll of the Senate in alphabetical order; each senator responds when his or her name is called. Senators who miss the roll call may still cast a vote as long as the recorded vote remains open. The vote is closed at the discretion of the presiding officer, but must remain open for a minimum of 15 minutes. If the vote is tied, the Vice President, if present, is entitled to a casting vote. If the Vice President is not present, however, the motion is resolved in the negative.
On occasion, the Senate may go into what is called a secret, or closed session. During a closed session, the chamber doors are closed, and the galleries are completely cleared of anyone not sworn to secrecy, not instructed in the rules of the closed session, or not essential to the session. Closed sessions are quite rare, and are usually held only under certain circumstances where the Senate is discussing sensitive subject-matter such as information critical to national security, private communications from the President, or even to discuss Senate deliberations during impeachment trials. Any Senator has the right to call a closed session as long as the motion is seconded.
Budget bills are governed under a special rule process called "Reconciliation" that disallows filibusters. Reconciliation was devised in 1974 but came into use in the early 1980s.
The Rules by number[edit]
The Standing Rules of the Senate detail the rules of order of the United States Senate. The latest version was adopted on April 27, 2000 and comprises the following 43 rules. The Legislative Transparency and Accountability Act of 2006 introduces a 44th rule on earmarks.
SR Rule I: Appointment of a Senator to the Chair
SR Rule II: Presentation of Credentials and Questions of Privilege
SR Rule III: Oaths
SR Rule IV: Commencement of Daily Sessions
SR Rule V: Suspension and Amendment of the Rules
SR Rule VI: Quorum - Absent Senators May Be Sent For
SR Rule VII: Morning Business
SR Rule VIII: Order of Business
SR Rule IX: Messages
SR Rule X: Special Orders
SR Rule XI: Papers - Withdrawal, Printing, Reading of, and Reference
SR Rule XII: Voting Procedure
SR Rule XIII: Reconsideration
SR Rule XIV: Joint Resolutions, and Preambles Thereto
SR Rule XV: Amendments and Resolutions
SR Rule XVI: Appropriations and Amendments to General Appropriation Bills
SR Rule XVII: Reference to Committees; Motions to Discharge; Reports of Committees; and Hearings Available
SR Rule XVIII: Business Continued from Session to Session
SR Rule XIX: Debate
SR Rule XX: Questions for Order
SR Rule XXI: Session with Closed Doors
SR Rule XXII: Precedence of Motions
SR Rule XXIII: Privilege of the Floor
SR Rule XXIV: Appointments of Committee
SR Rule XXV: Standing Committees
SR Rule XXVI: Committee Procedure
SR Rule XXVII: Committee Staff
SR Rule XXVIII: Conference Committees; Reports; Open Meetings
SR Rule XXIX: Executive Sessions
SR Rule XXX: Executive Session - Proceedings on Treaties
SR Rule XXXI: Executive Session - Proceedings on Nominations
SR Rule XXXII: The President Furnished with Copies of Record Executive Sessions
SR Rule XXXIII: Senate Chamber - Senate Wing of the Capitol
SR Rule XXXIV: Public Financial Disclosure
SR Rule XXXV: Gifts
SR Rule XXXVI: Outside Earned Income
SR Rule XXXVII: Conflict of Interest
SR Rule XXXVIII: Prohibition of Unofficial Office Accounts
SR Rule XXXIX: Foreign Travel
SR Rule XL: Franking Privilege and Radio and Television Studios
SR Rule XLI: Political Fund Activity; Definitions
SR Rule XLII: Employment Practices
SR Rule XLIII: Representation by Members
SR Rule XLIV: Congressionally Directed Spending and Related Items
References[edit]
Jump up ^ http://www.gpo.gov/fdsys/pkg/BILLS-113sres16ats/pdf/BILLS-113sres16ats.pdf
External links[edit]
Wikisource has original text related to this article:
Standing Rules of the Senate
Official text, Standing Rules of the Senate
Standing Rules of the Senate, One Hundred Thirteenth Congress, November 4, 2013
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