31:
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681:(or also heterotropic modulation) is a change in synaptic strength that results from the activity of other neurons. Again, the plasticity can alter the number of vesicles or their replenishment rate or the relationship between calcium and vesicle release. Additionally, it could directly affect calcium influx. Heterosynaptic plasticity can also be postsynaptic in nature, affecting receptor sensitivity.
560:, the number of release sites. "Unitary connection" usually refers to an unknown number of individual synapses connecting a presynaptic neuron to a postsynaptic neuron. The amplitude of postsynaptic potentials (PSPs) can be as low as 0.4 mV to as high as 20 mV. The amplitude of a PSP can be modulated by
312:
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Desensitization of the postsynaptic receptors is a decrease in response to the same neurotransmitter stimulus. It means that the strength of a synapse may in effect diminish as a train of action potentials arrive in rapid succession – a phenomenon that gives rise to the so-called frequency dependence
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farther away from the threshold, decreasing its excitability and making it more difficult for the neuron to initiate an action potential. If an IPSP overlaps with an EPSP, the IPSP can in many cases prevent the neuron from firing an action potential. In this way, the output of a neuron may depend on
726:
for eliciting an action potential. When action potentials from multiple presynaptic neurons fire simultaneously, or if a single presynaptic neuron fires at a high enough frequency, the EPSPs can overlap and summate. If enough EPSPs overlap, the summated EPSP can reach the threshold for initiating an
275:
Synapses may be described as symmetric or asymmetric. When examined under an electron microscope, asymmetric synapses are characterized by rounded vesicles in the presynaptic cell, and a prominent postsynaptic density. Asymmetric synapses are typically excitatory. Symmetric synapses in contrast have
776:, assigning a role for volume transmission in the control of ionic and neurotransmitter homeostasis. Approximately 78% of neurogliaform cell boutons do not form classical synapses. This may be the first definitive example of neurons communicating chemically where classical synapses are not present.
599:
Synaptic transmission can be changed by previous activity. These changes are called synaptic plasticity and may result in either a decrease in the efficacy of the synapse, called depression, or an increase in efficacy, called potentiation. These changes can either be long-term or short-term. Forms
759:
When a neurotransmitter is released at a synapse, it reaches its highest concentration inside the narrow space of the synaptic cleft, but some of it is certain to diffuse away before being reabsorbed or broken down. If it diffuses away, it has the potential to activate receptors that are located
307:
Here is a summary of the sequence of events that take place in synaptic transmission from a presynaptic neuron to a postsynaptic cell. Each step is explained in more detail below. Note that with the exception of the final step, the entire process may run only a few hundred microseconds, in the
808:
of each other, rather than the 20 to 40 nm distance that separates cells at chemical synapses. As opposed to chemical synapses, the postsynaptic potential in electrical synapses is not caused by the opening of ion channels by chemical transmitters, but rather by direct electrical coupling
352:
These proteins change shape, causing the membranes of some "docked" vesicles to fuse with the membrane of the presynaptic cell, thereby opening the vesicles and dumping their neurotransmitter contents into the synaptic cleft, the narrow space between the membranes of the pre- and postsynaptic
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Romanov, Roman A.; Lasher, Robert S.; High, Brigit; Savidge, Logan E.; Lawson, Adam; Rogachevskaja, Olga A.; Zhao, Haitian; Rogachevsky, Vadim V.; Bystrova, Marina F.; Churbanov, Gleb D.; Adameyko, Igor; Harkany, Tibor; Yang, Ruibiao; Kidd, Grahame J.; Marambaud, Philippe; Kinnamon, John C.;
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currents. Whether a synapse is excitatory or inhibitory depends on what type(s) of ion channel conduct the postsynaptic current(s), which in turn is a function of the type of receptors and neurotransmitter employed at the synapse. The second way a receptor can affect membrane potential is by
645:. Homosynaptic plasticity can affect the number and replenishment rate of vesicles or it can affect the relationship between calcium and vesicle release. Homosynaptic plasticity can also be postsynaptic in nature. It can result in either an increase or decrease in synaptic strength.
825:. While chemical synapses are found between both excitatory and inhibitory neurons, electrical synapses are most commonly found between smaller local inhibitory neurons. Electrical synapses can exist between two axons, two dendrites, or between an axon and a dendrite. In some
1924:
855:. Synapses are affected by drugs, such as curare, strychnine, cocaine, morphine, alcohol, LSD, and countless others. These drugs have different effects on synaptic function, and often are restricted to synapses that use a specific neurotransmitter. For example,
641:(or also homotropic modulation) is a change in the synaptic strength that results from the history of activity at a particular synapse. This can result from changes in presynaptic calcium as well as feedback onto presynaptic receptors, i.e. a form of
313:
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Proteins in the PSD are involved in anchoring and trafficking neurotransmitter receptors and modulating the activity of these receptors. The receptors and PSDs are often found in specialized protrusions from the main dendritic shaft called
201:
Synapses are functional connections between neurons, or between neurons and other types of cells. A typical neuron gives rise to several thousand synapses, although there are some types that make far fewer. Most synapses connect
771:
can inhibit other nearby cortical neurons by releasing the neurotransmitter GABA into the extracellular space. Along the same vein, GABA released from neurogliaform cells into the extracellular space also acts on surrounding
316:
260:; for synapses between two neurons the postsynaptic region may be found on the dendrites or cell body. Immediately behind the postsynaptic membrane is an elaborate complex of interlinked proteins called the
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Craig C. Garner and Kang Shen. Structure and
Function of Vertebrate and Invertebrate Active Zones. Structure and Functional Organization of the Synapse. Ed: Johannes Hell and Michael Ehlers. Springer, 2008.
367:
in some way. Several types of activation are possible, as described in more detail below. In any case, this is the key step by which the synaptic process affects the behavior of the postsynaptic cell.
438:. As a whole, the protein complex or structure that mediates the docking and fusion of presynaptic vesicles is called the active zone. The membrane added by the fusion process is later retrieved by
522:
The neurotransmitter may diffuse away due to thermally-induced oscillations of both it and the receptor, making it available to be broken down metabolically outside the neuron or to be reabsorbed.
287:—is a gap between the pre- and postsynaptic cells that is about 20 nm (0.02 μ) wide. The small volume of the cleft allows neurotransmitter concentration to be raised and lowered rapidly.
767:
Recent work indicates that volume transmission may be the predominant mode of interaction for some special types of neurons. In the mammalian cerebral cortex, a class of neurons called
2539:
2084:
Bennett MV, Pappas GD, Aljure E, Nakajima Y (March 1967). "Physiology and ultrastructure of electrotonic junctions. II. Spinal and medullary electromotor nuclei in mormyrid fish".
50:
molecules. The neurotransmitter binds to chemical receptor molecules located in the membrane of another neuron, the postsynaptic neuron, on the opposite side of the synaptic cleft.
2443:
Hormuzdi, SG; Filippov, MA; Mitropoulou, G; Monyer, H; Bruzzone, R (March 2004). "Electrical synapses: a dynamic signaling system that shapes the activity of neuronal networks".
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Receptors on the opposite side of the synaptic gap bind neurotransmitter molecules. Receptors can respond in either of two general ways. First, the receptors may directly open
374:, the motion of atoms, vibrating about their equilibrium positions in a crystalline solid, neurotransmitter molecules eventually break loose from the receptors and drift away.
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1805:"From the Golgi-Cajal mapping to the transmitter-based characterization of the neuronal networks leading to two modes of brain communication: wiring and volume transmission"
225:
Chemical synapses pass information directionally from a presynaptic cell to a postsynaptic cell and are therefore asymmetric in structure and function. The presynaptic
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the input of many different neurons, each of which may have a different degree of influence, depending on the strength and type of synapse with that neuron.
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between both neurons. Electrical synapses are faster than chemical synapses. Electrical synapses are found throughout the nervous system, including in the
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After a neurotransmitter molecule binds to a receptor molecule, it must be removed to allow for the postsynaptic membrane to continue to relay subsequent
957:'I felt the need of some name to call the junction between nerve-cell and nerve-cell... I suggested using "syndesm"... He consulted his Trinity friend
418:
proteins found within the membranes of the synaptic vesicles, allowing the vesicles to fuse with the presynaptic membrane. The fusion of a vesicle is a
1648:"Target-specific regulation of synaptic efficacy in the feeding central pattern generator of Aplysia: potential substrates for behavioral plasticity?"
931:. Based on these observations, they developed the 'quantal hypothesis' that is the basis for our current understanding of neurotransmitter release as
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The neurotransmitter is either reabsorbed by the presynaptic cell, and then repackaged for future release, or else it is broken down metabolically.
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or can change as a result of previous activity. Changes in the synaptic strength can be short-term, lasting seconds to minutes, or long-term (
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568:, or LTP), lasting hours. Learning and memory are believed to result from long-term changes in synaptic strength, via a mechanism known as
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Axons connecting dendrite to dendrite are dendrodendritic synapses. Axons which connect axon to dendrite are called axodendritic synapses
764:. It is well established that such effects occur to some degree, but their functional importance has long been a matter of controversy.
1342:
1242:
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An exception to the general trend of neurotransmitter release by vesicular fusion is found in the type II receptor cells of mammalian
2119:
Pereda AE, Rash JE, Nagy JI, Bennett MV (December 2004). "Dynamics of electrical transmission at club endings on the
Mauthner cells".
624:. Synaptic plasticity can be either homosynaptic (occurring at a single synapse) or heterosynaptic (occurring at multiple synapses).
502:
inside the postsynaptic neuron. These second messengers can then amplify the inhibitory or excitatory response to neurotransmitters.
46:. When the action potential reaches the presynaptic terminal, it provokes the release of a synaptic vesicle, secreting its quanta of
1579:"Chemical synapses without synaptic vesicles: Purinergic neurotransmission through a CALHM1 channel-mitochondrial signaling complex"
2619:
581:
of synapses. The nervous system exploits this property for computational purposes, and can tune its synapses through such means as
276:
flattened or elongated vesicles, and do not contain a prominent postsynaptic density. Symmetric synapses are typically inhibitory.
218:
except as points where the membranes of two cells appear to touch, but their cellular elements can be visualized clearly using an
760:
either at other synapses or on the membrane away from any synapse. The extrasynaptic activity of a neurotransmitter is known as
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Axons terminating on the postsynaptic cell body are axosomatic synapses. Axons that terminate on axons are axoaxonic synapses
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containing neurotransmitter are localized near the synaptic membrane. The arriving action potential produces an influx of
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coined the word 'synapse' and the history of the word was given by
Sherrington in a letter he wrote to John Fulton:
3243:
3173:
875:, which causes the body to pick up and react to weaker and previously ignored stimuli, resulting in uncontrollable
34:
Artistic interpretation of the major elements in chemical synaptic transmission. An electrochemical wave called an
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An electron microscope picture gallery assembled by
Kristen Harris' lab of synapses and other neuronal structures.
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process, leading to frequent failure of synaptic transmission at the very small synapses that are typical for the
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Calcium ions flow through the presynaptic membrane, rapidly increasing the calcium concentration in the interior.
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One of the most important features of chemical synapses is that they are the site of action for the majority of
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on the postsynaptic cell. Finally, the neurotransmitters are cleared from the synapse through one of several
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is a chemical (or electrical) synapse formed when the axon of one neuron synapses with its own dendrites.
2893:
2831:
1925:"Unitary GABAergic volume transmission from individual interneurons to astrocytes in the cerebral cortex"
705:
494:
2888:
1729:
Bruce
Alberts; Alexander Johnson; Julian Lewis; Martin Raff; Keith Roberts; Peter Walter, eds. (2002).
965:
scholar, about it, and
Verrall suggested "synapse" (from the Greek "clasp").'–Charles Scott Sherrington
727:
action potential. This process is known as summation, and can serve as a high pass filter for neurons.
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in the presynaptic neuron will trigger an action potential in the postsynaptic cell. In many cases the
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performed some of the important early experiments on synaptic integration, for which he received the
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2375:"Relationship between presynaptic calcium current and postsynaptic potential in squid giant synapse"
2133:
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Zoli M, Torri C, Ferrari R, et al. (1998). "The emergence of the volume transmission concept".
1461:"Relationship between presynaptic calcium current and postsynaptic potential in squid giant synapse"
3178:
2904:
1445:
1375:
Widrow, Bernard; Kim, Youngsik; Park, Dookun; Perin, Jose Krause (2019). "Nature's
Learning Rule".
1127:
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107:) that is adjacent to another neuron. The neurotransmitters are contained within small sacs called
24:
556:(the postsynaptic response to the release of a single neurotransmitter vesicle, a 'quantum'), and
229:, or synaptic bouton, is a specialized area within the axon of the presynaptic cell that contains
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in the postsynaptic cell membrane, causing ions to enter or exit the cell and changing the local
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is estimated to contain from 10 to 5 × 10 (100–500 trillion) synapses. Every cubic millimeter of
692:, which, in addition, generates an inhibitory effect on presynaptic terminals of neurons of the
210:, but there are also other types of connections, including axon-to-cell-body, axon-to-axon, and
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and Katz advanced the hypothesis that depolarization-induced influx of calcium ions triggers
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The neurotransmitter diffuses within the cleft. Some of it escapes, but some of it binds to
242:
176:
153:
2037:"Electrical synapses mediate signal transmission in the rod pathway of the mammalian retina"
1437:
1244:
We're Born to Learn: Using the Brain's
Natural Learning Process to Create Today's Curriculum
457:
is released directly from the cytoplasm into the synaptic cleft via voltage gated channels.
19:
This article is about chemical synapses of the nervous system. For general information, see
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On the other hand, a presynaptic neuron releasing an inhibitory neurotransmitter, such as
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of the membrane at the synapse causes channels to open that are permeable to calcium ions.
179:
also exist. Without a qualifier, however, "synapse" commonly refers to chemical synapses.
8:
3368:
3213:
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2174:"Bernard Katz, quantal transmitter release and the foundations of presynaptic physiology"
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833:, electrical synapses can be found within the same terminal of a chemical synapse, as in
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The high calcium concentration activates a set of calcium-sensitive proteins attached to
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Enzymes within the subsynaptic membrane may inactivate/metabolize the neurotransmitter.
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Immediately opposite is a region of the postsynaptic cell containing neurotransmitter
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at the down stroke of the action potential (tail current). Calcium ions then bind to
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The release of a neurotransmitter is triggered by the arrival of a nerve impulse (or
357:
1959:
1789:
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1553:
1516:
Chapman, Edwin R. (2002). "Synaptotagmin: A Ca2+ sensor that triggers exocytosis?".
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1038:
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2009:
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1868:"Regulation of cortical microcircuits by unitary GABA-mediated volume transmission"
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Alonso-Nanclares L, Gonzalez-Soriano J, Rodriguez JR, DeFelipe J (September 2008).
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traveling along the membrane of the presynaptic cell, until it reaches the synapse.
325:
234:
230:
215:
120:
108:
97:
47:
35:
2322:"Transmission by presynaptic spike-like depolarization in the squid giant synapse"
434:
is driven by the action of a set of proteins in the presynaptic terminal known as
30:
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90:
61:' signals can be sent to each other and to non-neuronal cells such as those in
1943:
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2728:
2097:
1602:
1537:
958:
876:
860:
689:
653:
532:
415:
226:
1078:
660:, inhibiting further release of noradrenaline. This effect is utilized with
656:, which, besides affecting postsynaptic receptors, also affects presynaptic
430:), on the other hand, have a synaptic release probability, in effect, of 1.
237:(as well as a number of other supporting structures and organelles, such as
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2733:
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Rózsa M, Baka J, Bordé S, Rózsa B, Katona G, Tamás G, et al. (2015).
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1502:
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Artificial
Intelligence in the Age of Neural Networks and Brain Computing
822:
439:
250:
2557:
was created from a revision of this article dated 19 June 2005
1891:
1636:
Sherwood L., stikawy (2007). Human
Physiology 6e: From Cells to Systems
1340:
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2959:
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1671:
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932:
868:
531:
pumps may actively pump the neurotransmitter back into the presynaptic
419:
399:
398:) and occurs through an unusually rapid process of cellular secretion (
324:
The process begins with a wave of electrochemical excitation called an
112:
82:
1832:
3408:
2942:
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962:
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1997:
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for reprocessing and re-release following a later action potential.
528:
363:
The binding of neurotransmitter causes the receptor molecule to be
207:
124:
100:
1126:
Cowan, W. Maxwell; SĂĽdhof, Thomas C.; Stevens, Charles F. (2003).
3335:
3296:
3188:
2597:
2495:
Nicholls, J.G.; Martin, A.R.; Wallace, B.G.; Fuchs, P.A. (2001).
2442:
1984:
1731:"Ch. 11. Section: Single Neurons Are Complex Computation Devices"
1274:
From Computer to Brain: Foundations of Computational Neuroscience
1186:
904:
872:
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that is formed at a narrow gap between the pre- and postsynaptic
474:
390:
Release of neurotransmitter occurs at the end of axonal branches.
291:
187:
168:
160:
has separately been estimated at 0.15 quadrillion (150 trillion)
86:
20:
1703:
1444:. Philadelphia, PA: Lippincott Williams & Wilkins. pp.
2995:
2950:
2715:
2424:
Bear, Mark F.; Connors, Barry W.; Paradiso, Michael A. (2001).
1708:(5th ed.). Edinburgh: Churchill Livingstone. p. 129.
856:
810:
793:
167:
in 1897. Chemical synapses are not the only type of biological
58:
43:
16:
Biological junctions through which neurons' signals can be sent
214:. Synapses are generally too small to be recognizable using a
927:
observed spontaneous miniature synaptic currents at the frog
446:
for the formation of fresh neurotransmitter-filled vesicles.
145:
142:
139:
66:
1436:
Bear, Mark F; Connors, Barry W; Paradiso, Michael A (2007).
1341:
Weiss, Mirin; Dr Steven M. Mirin; Dr Roxanne Bartel (1994).
3034:
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892:
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molecules located on the membrane of the postsynaptic cell.
203:
39:
2494:
1213:
896:
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increases the inhibitory effects of the neurotransmitter
804:. At gap junctions, cells approach within about 3.5
518:. This removal can happen through one or more processes:
156:, i.e. 10) of them. The number of synapses in the human
1575:
1187:
Squire, Larry R.; Floyd Bloom; Nicholas Spitzer (2008).
1054:"Gender differences in human cortical synaptic density"
899:
interferes with synapses that use the neurotransmitter
871:
blocks the inhibitory effects of the neurotransmitter
779:
2428:. Hagerstown, MD: Lippincott Williams & Wilkins.
2372:
2295:
2118:
1972:
1458:
1418:
863:
from depolarizing the postsynaptic membrane, causing
93:
to connect to and control other systems of the body.
2501:(4th ed.). Sunderland, MA: Sinauer Associates.
2480:(4th ed.). Hoboken, NJ: John Wiley & Sons.
2476:
Cell and Molecular Biology: concepts and experiments
1803:
Fuxe K, Dahlström A, Höistad M, et al. (2007).
1802:
1577:
Kolesnikov, Stanislav S.; Finger, Thomas E. (2018).
548:
as the product of (presynaptic) release probability
2423:
2319:
1922:
245:). Synaptic vesicles are docked at the presynaptic
2473:
1374:
1303:
1125:
195:Further information on formation of synapses:
2299:; Schwartz, James H.; Jessell, Thomas M. (2000).
2172:Augustine, George J.; Kasai, Haruo (2007-02-01).
1569:
1009:Drachman D (2005). "Do we have brain to spare?".
699:
412:voltage-dependent, calcium-selective ion channels
297:
135:to terminate the action of the neurotransmitter.
3430:
1759:
1214:Hyman, Steven E.; Eric Jonathan Nestler (1993).
738:(IPSP) in the postsynaptic neuron, bringing the
233:enclosed in small membrane-bound spheres called
131:either on the presynaptic cell or on some other
2281:(9th ed.). Boston, MA: Pearson Education.
1866:Oláh S, Füle M, Komlósi G, et al. (2009).
1865:
1645:
2171:
2034:
544:The strength of a synapse has been defined by
111:, and are released into the synaptic cleft by
2613:
2320:Llinás R, Sugimori M, Simon SM (April 1982).
1704:Rang, H.P.; Dale, M.M.; Ritter, J.M. (2003).
1414:
1412:
1166:. W. W. Norton & Company. pp. 1–37.
1998:"Electrical synapses in the mammalian brain"
1220:. American Psychiatric Pub. pp. 425–6.
667:
1995:
1861:
1859:
1632:
1630:
1163:Nerve Endings: The Discovery of the Synapse
575:
381:
96:At a chemical synapse, one neuron releases
2620:
2606:
1409:
1240:
1008:
664:to perform inhibitory effects on the SNS.
627:
402:). Within the presynaptic nerve terminal,
69:. Chemical synapses allow neurons to form
2406:
2373:Llinás R, Steinberg IZ, Walton K (1981).
2355:
2345:
2245:
2197:
2132:
2060:
1899:
1831:
1610:
1492:
1459:Llinás R, Steinberg IZ, Walton K (1981).
1297:
1295:
1087:
1077:
191:Diagram of a chemical synaptic connection
163:The word "synapse" was introduced by Sir
2565:, and does not reflect subsequent edits.
2548:
1856:
1627:
1452:
1347:. American Psychiatric Pub. p. 52.
1336:
1334:
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1264:
914:
385:
186:
29:
2305:(4th ed.). New York: McGraw-Hill.
2271:
2035:Veruki ML, Hartveit E (December 2002).
1699:
1697:
1515:
1217:The Molecular Foundations of Psychiatry
1159:
1121:
1119:
57:are biological junctions through which
3431:
2014:10.1146/annurev.neuro.26.041002.131128
1301:
1292:
1270:
754:
749:Nobel Prize for Physiology or Medicine
588:
2601:
2592:Atlas of Ultrastructure Neurocytology
1518:Nature Reviews Molecular Cell Biology
1431:
1429:
1427:
1331:
1261:
937:Nobel Prize in Physiology or Medicine
911:and therefore increases its effects.
2471:
1694:
1646:DĂaz-RĂos M, Miller MW (June 2006).
1326:synapses connect axons to cell body.
1116:
684:One example is again neurons of the
539:
426:. Large chemical synapses (e.g. the
1973:Kandel, Schwartz & Jessell 2000
1419:Kandel, Schwartz & Jessell 2000
840:
780:Relationship to electrical synapses
460:
123:including enzymatic degradation or
13:
2627:
2535:
2053:10.1523/JNEUROSCI.22-24-10558.2002
1424:
1385:10.1016/b978-0-12-815480-9.00001-3
1193:. Academic Press. pp. 425–6.
1023:10.1212/01.WNL.0000166914.38327.BB
310:
14:
3460:
2516:
2426:Neuroscience: Exploring the Brain
2143:10.1016/j.brainresrev.2004.06.010
1824:10.1016/j.brainresrev.2007.02.009
1739:(4th ed.). Garland Science.
1440:Neuroscience: exploring the brain
815:reticular nucleus of the thalamus
736:inhibitory postsynaptic potential
720:excitatory postsynaptic potential
2587:Synapse – Cell Centered Database
2547:
935:and for which Katz received the
2784:Oligodendrocyte progenitor cell
2214:
2165:
2112:
2077:
2028:
1989:
1978:
1966:
1916:
1796:
1753:
1722:
1639:
1560:
1509:
1310:. John Wiley and Sons. p.
951:Sir Charles Scott Sherringtonin
115:. These molecules then bind to
1306:Biology: A Self-Teaching Guide
1234:
1207:
1180:
1153:
1104:
1045:
1002:
700:Integration of synaptic inputs
694:parasympathetic nervous system
648:One example is neurons of the
505:
298:Signaling in chemical synapses
104:
1:
2399:10.1016/S0006-3495(81)84899-0
2265:
1774:10.1016/S0165-0173(97)00048-9
1736:Molecular Biology of the Cell
1485:10.1016/S0006-3495(81)84899-0
498:modulating the production of
2457:10.1016/j.bbamem.2003.10.023
2326:Proc. Natl. Acad. Sci. U.S.A
2302:Principles of Neural Science
2190:10.1113/jphysiol.2006.123224
1996:Connors BW, Long MA (2004).
1932:Brain Structure and Function
1247:. Corwin Press. p. 56.
1160:Rapport, Richard L. (2005).
1058:Proc. Natl. Acad. Sci. U.S.A
939:in 1970. In the late 1960s,
481:. In general, the result is
453:. Here the neurotransmitter
182:
152:contains roughly a billion (
7:
2894:Postganglionic nerve fibers
1379:. Elsevier. pp. 1–30.
1271:Lytton, William W. (2002).
969:
706:Summation (neurophysiology)
302:
10:
3465:
2889:Preganglionic nerve fibers
1302:Garber, Steven D. (2002).
883:acts on synapses that use
844:
792:link between two abutting
722:(EPSP) will not reach the
703:
686:sympathetic nervous system
671:
650:sympathetic nervous system
631:
592:
585:of the proteins involved.
473:. The resulting change in
194:
117:neurotransmitter receptors
77:. They are crucial to the
18:
3399:Olfactory receptor neuron
3354:
3295:
3288:
3224:
3154:
3111:
3071:
3063:Neurofibril/neurofilament
3033:
3015:
3008:
2994:
2941:
2913:
2819:
2810:
2757:
2714:
2707:
2644:
2635:
2232:(5729): 190. 1970-10-24.
2178:The Journal of Physiology
2121:Brain Res. Brain Res. Rev
1944:10.1007/s00429-015-1166-9
1762:Brain Res. Brain Res. Rev
1595:10.1126/scisignal.aao1815
1241:Smilkstein, Rita (2003).
1132:. JHU Press. p. 11.
976:Acclimatisation (neurons)
679:Heterosynaptic plasticity
674:Heterosynaptic plasticity
668:Heterosynaptic plasticity
467:ligand-gated ion channels
165:Charles Scott Sherrington
2098:10.1152/jn.1967.30.2.180
1277:. Springer. p. 28.
1190:Fundamental Neuroscience
996:
576:Receptor desensitization
382:Neurotransmitter release
103:into a small space (the
25:synapse (disambiguation)
2582:Synapse Review for Kids
2226:British Medical Journal
1111:Brain Facts and Figures
1079:10.1073/pnas.0803652105
887:neurotransmitters, and
859:is a poison that stops
658:α2-adrenergic receptors
639:Homosynaptic plasticity
634:Homosynaptic plasticity
628:Homosynaptic plasticity
471:transmembrane potential
79:biological computations
3346:Neuromuscular junction
3209:III or Aδ or fast pain
2543:
2523:Listen to this article
2347:10.1073/pnas.79.7.2415
2278:Physiology of Behavior
2238:10.1136/bmj.4.5729.190
1113:Washington University.
967:
929:neuromuscular junction
622:long-term potentiation
566:long-term potentiation
479:postsynaptic potential
428:neuromuscular junction
424:central nervous system
391:
320:
192:
177:immunological synapses
75:central nervous system
51:
23:. For other uses, see
2542:
2472:Karp, Gerald (2005).
955:
915:History and etymology
714:is strong enough, an
652:(SNS), which release
610:synaptic augmentation
602:short-term plasticity
389:
319:
243:endoplasmic reticulum
190:
129:specific transporters
33:
3364:Meissner's corpuscle
3329:Postsynaptic density
3226:Efferent nerve fiber
3214:IV or C or slow pain
3156:Afferent nerve fiber
2982:Satellite glial cell
2574:More spoken articles
2498:From Neuron to Brain
2445:Biochim Biophys Acta
1985:Hormuzdi et al. 2004
618:long-term depression
614:long-term plasticity
262:postsynaptic density
212:dendrite-to-dendrite
121:potential mechanisms
3444:Signal transduction
3369:Merkel nerve ending
2391:1981BpJ....33..323L
2379:Biophysical Journal
2338:1982PNAS...79.2415L
2002:Annu. Rev. Neurosci
1892:10.1038/nature08503
1884:2009Natur.461.1278O
1477:1981BpJ....33..323L
1465:Biophysical Journal
1070:2008PNAS..10514615A
907:blocks reuptake of
788:is an electrically
769:neurogliaform cells
762:volume transmission
755:Volume transmission
643:autocrine signaling
595:Synaptic plasticity
589:Synaptic plasticity
570:synaptic plasticity
500:chemical messengers
220:electron microscope
3404:Photoreceptor cell
3374:Pacinian corpuscle
3305:Electrical synapse
3259:Lower motor neuron
3254:Upper motor neuron
2975:Internodal segment
2915:Connective tissues
2885:Autonomic ganglion
2544:
919:During the 1950s,
853:psychoactive drugs
786:electrical synapse
740:membrane potential
712:excitatory synapse
710:In general, if an
608:or depression and
392:
321:
308:fastest synapses.
249:at regions called
193:
52:
38:travels along the
3426:
3425:
3422:
3421:
3389:Free nerve ending
3356:Sensory receptors
3284:
3283:
3199:Ib or Golgi or Aα
3107:
3106:
2990:
2989:
2867:Ramus communicans
2806:
2805:
2802:
2801:
2672:Commissural fiber
2667:Association fiber
2662:Projection fibers
2540:
2508:978-0-87893-439-3
2487:978-0-471-46580-5
2435:978-0-7817-3944-3
2312:978-0-8385-7701-1
2288:978-0-205-59389-7
2184:(Pt 3): 623–625.
1878:(7268): 1278–81.
1746:978-0-8153-3218-3
1715:978-0-443-07145-4
1589:(529): eaao1815.
1583:Science Signaling
1394:978-0-12-815480-9
1354:978-1-58562-138-5
1321:978-0-471-22330-6
1284:978-0-387-95526-1
1254:978-0-7619-4642-7
1227:978-0-88048-353-7
1200:978-0-12-374019-9
1173:978-0-393-06019-5
847:Neuropharmacology
745:John Carew Eccles
540:Synaptic strength
372:thermal vibration
358:chemical receptor
317:
235:synaptic vesicles
231:neurotransmitters
109:synaptic vesicles
89:. They allow the
55:Chemical synapses
3456:
3319:Synaptic vesicle
3314:Chemical synapse
3293:
3292:
3013:
3012:
3006:
3005:
2817:
2816:
2712:
2711:
2642:
2641:
2622:
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2608:
2599:
2598:
2564:
2562:
2551:
2550:
2541:
2531:
2529:
2524:
2512:
2491:
2479:
2468:
2451:(1–2): 113–137.
2439:
2420:
2410:
2369:
2359:
2349:
2316:
2292:
2273:Carlson, Neil R.
2260:
2259:
2249:
2218:
2212:
2211:
2201:
2169:
2163:
2162:
2136:
2116:
2110:
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2075:
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2064:
2047:(24): 10558–66.
2032:
2026:
2025:
1993:
1987:
1982:
1976:
1970:
1964:
1963:
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1920:
1914:
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1863:
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1108:
1102:
1101:
1091:
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1042:
1006:
841:Effects of drugs
716:action potential
688:, which release
606:synaptic fatigue
461:Receptor binding
396:action potential
347:neurotransmitter
326:action potential
318:
270:dendritic spines
216:light microscope
98:neurotransmitter
48:neurotransmitter
36:action potential
3464:
3463:
3459:
3458:
3457:
3455:
3454:
3453:
3429:
3428:
3427:
3418:
3350:
3280:
3229:
3220:
3204:II or Aβ and Aγ
3159:
3150:
3103:
3093:Apical dendrite
3088:Dendritic spine
3067:
3029:
2999:
2986:
2970:Node of Ranvier
2965:Myelin incisure
2937:
2909:
2798:
2789:Oligodendrocyte
2772:Ependymal cells
2753:
2703:
2631:
2626:
2578:
2577:
2566:
2560:
2558:
2555:This audio file
2552:
2545:
2536:
2533:
2527:
2526:
2522:
2519:
2509:
2488:
2436:
2313:
2297:Kandel, Eric R.
2289:
2268:
2263:
2220:
2219:
2215:
2170:
2166:
2134:10.1.1.662.9352
2127:(1–3): 227–44.
2117:
2113:
2086:J. Neurophysiol
2082:
2078:
2033:
2029:
1994:
1990:
1983:
1979:
1971:
1967:
1927:
1921:
1917:
1864:
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1797:
1768:(2–3): 136–47.
1758:
1754:
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1727:
1723:
1716:
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1695:
1664:10.2307/4134559
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734:, can cause an
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583:phosphorylation
578:
562:neuromodulators
552:, quantal size
542:
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495:hyperpolarizing
493:in the case of
485:in the case of
463:
384:
345:that contain a
331:The electrical
311:
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300:
247:plasma membrane
199:
185:
158:cerebral cortex
150:cerebral cortex
28:
17:
12:
11:
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3462:
3452:
3451:
3449:Neural synapse
3446:
3441:
3439:Cell signaling
3424:
3423:
3420:
3419:
3417:
3416:
3414:Taste receptor
3411:
3406:
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3396:
3391:
3386:
3384:Muscle spindle
3381:
3379:Ruffini ending
3376:
3371:
3366:
3360:
3358:
3352:
3351:
3349:
3348:
3343:
3341:Ribbon synapse
3338:
3333:
3332:
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3326:
3321:
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3301:
3299:
3290:
3286:
3285:
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3161:Sensory neuron
3152:
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3129:Pseudounipolar
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3099:
3097:Basal dendrite
3090:
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3055:
3050:
3048:Axon terminals
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2629:Nervous tissue
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2517:External links
2515:
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2385:(3): 323–352.
2370:
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2008:(1): 393–418.
1988:
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1938:(1): 651–659.
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1530:10.1038/nrm855
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1017:(12): 2004–5.
1000:
998:
995:
994:
993:
991:Ribbon synapse
988:
983:
978:
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968:
941:Ricardo Miledi
916:
913:
845:Main article:
842:
839:
835:Mauthner cells
781:
778:
756:
753:
704:Main article:
701:
698:
672:Main article:
669:
666:
632:Main article:
629:
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593:Main article:
590:
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489:currents, and
462:
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432:Vesicle fusion
383:
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368:
361:
354:
350:
339:
336:
333:depolarization
329:
304:
301:
299:
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281:synaptic cleft
197:Synaptogenesis
184:
181:
105:synaptic cleft
91:nervous system
81:that underlie
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3274:Îł motorneuron
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3269:β motorneuron
3267:
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3264:α motorneuron
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2380:
2376:
2371:
2367:
2363:
2358:
2353:
2348:
2343:
2339:
2335:
2332:(7): 2415–9.
2331:
2327:
2323:
2318:
2314:
2308:
2304:
2303:
2298:
2294:
2290:
2284:
2280:
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2270:
2269:
2257:
2253:
2248:
2243:
2239:
2235:
2231:
2227:
2223:
2222:"Nobel prize"
2217:
2209:
2205:
2200:
2195:
2191:
2187:
2183:
2179:
2175:
2168:
2160:
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2103:
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2095:
2091:
2087:
2080:
2072:
2068:
2063:
2058:
2054:
2050:
2046:
2042:
2038:
2031:
2023:
2019:
2015:
2011:
2007:
2003:
1999:
1992:
1986:
1981:
1975:, p. 176
1974:
1969:
1961:
1957:
1953:
1949:
1945:
1941:
1937:
1933:
1926:
1919:
1911:
1907:
1902:
1897:
1893:
1889:
1885:
1881:
1877:
1873:
1869:
1862:
1860:
1851:
1847:
1843:
1839:
1834:
1829:
1825:
1821:
1817:
1813:
1812:Brain Res Rev
1806:
1799:
1791:
1787:
1783:
1779:
1775:
1771:
1767:
1763:
1756:
1748:
1742:
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1737:
1732:
1725:
1717:
1711:
1707:
1700:
1698:
1689:
1685:
1681:
1677:
1673:
1669:
1665:
1661:
1658:(3): 215–29.
1657:
1653:
1649:
1642:
1633:
1631:
1622:
1618:
1613:
1608:
1604:
1600:
1596:
1592:
1588:
1584:
1580:
1572:
1563:
1555:
1551:
1547:
1543:
1539:
1535:
1531:
1527:
1523:
1519:
1512:
1504:
1500:
1495:
1490:
1486:
1482:
1478:
1474:
1470:
1466:
1462:
1455:
1447:
1442:
1441:
1432:
1430:
1428:
1421:, p. 182
1420:
1415:
1413:
1404:
1400:
1396:
1390:
1386:
1382:
1378:
1371:
1356:
1350:
1346:
1345:
1337:
1335:
1327:
1323:
1317:
1313:
1308:
1307:
1298:
1296:
1286:
1280:
1276:
1275:
1267:
1265:
1256:
1250:
1246:
1245:
1237:
1229:
1223:
1219:
1218:
1210:
1202:
1196:
1192:
1191:
1183:
1175:
1169:
1165:
1164:
1156:
1141:
1139:9780801871184
1135:
1131:
1130:
1122:
1120:
1112:
1107:
1099:
1095:
1090:
1085:
1080:
1075:
1071:
1067:
1063:
1059:
1055:
1048:
1040:
1036:
1032:
1028:
1024:
1020:
1016:
1012:
1005:
1001:
992:
989:
987:
984:
982:
979:
977:
974:
973:
966:
964:
960:
954:
952:
948:
946:
942:
938:
934:
930:
926:
922:
912:
910:
906:
902:
898:
894:
890:
886:
882:
878:
877:muscle spasms
874:
870:
866:
862:
861:acetylcholine
858:
854:
848:
838:
836:
832:
828:
824:
821:, and in the
820:
816:
812:
807:
803:
800:, known as a
799:
795:
791:
787:
777:
775:
770:
765:
763:
752:
750:
746:
741:
737:
733:
728:
725:
721:
717:
713:
707:
697:
695:
691:
690:noradrenaline
687:
682:
680:
675:
665:
663:
659:
655:
654:noradrenaline
651:
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640:
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623:
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607:
603:
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559:
555:
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547:
534:
533:axon terminal
530:
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517:
513:
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501:
496:
492:
488:
484:
480:
476:
472:
468:
458:
456:
452:
447:
445:
441:
437:
433:
429:
425:
421:
417:
416:synaptotagmin
413:
409:
405:
401:
397:
388:
376:
373:
369:
366:
362:
359:
355:
351:
348:
344:
340:
337:
334:
330:
327:
323:
322:
309:
295:
293:
288:
286:
283:—also called
282:
277:
273:
271:
265:
263:
259:
254:
252:
248:
244:
240:
236:
232:
228:
227:axon terminal
223:
221:
217:
213:
209:
205:
198:
189:
180:
178:
174:
170:
166:
161:
159:
155:
151:
147:
144:
141:
136:
134:
130:
126:
122:
118:
114:
110:
106:
102:
99:
94:
92:
88:
84:
80:
76:
72:
68:
64:
60:
56:
49:
45:
41:
37:
32:
26:
22:
3313:
3309:Gap junction
3231:Motor neuron
3025:Axon hillock
3001:nerve fibers
2955:Schwann cell
2865:
2848:
2826:
2744:Medium spiny
2657:White matter
2645:Tissue Types
2497:
2475:
2448:
2444:
2425:
2382:
2378:
2329:
2325:
2301:
2277:
2229:
2225:
2216:
2181:
2177:
2167:
2124:
2120:
2114:
2089:
2085:
2079:
2044:
2040:
2030:
2005:
2001:
1991:
1980:
1968:
1935:
1931:
1918:
1875:
1871:
1818:(1): 17–54.
1815:
1811:
1798:
1765:
1761:
1755:
1735:
1724:
1706:Pharmacology
1705:
1655:
1651:
1641:
1586:
1582:
1571:
1562:
1521:
1517:
1511:
1468:
1464:
1454:
1439:
1376:
1370:
1358:. Retrieved
1343:
1325:
1305:
1273:
1243:
1236:
1216:
1209:
1189:
1182:
1162:
1155:
1143:. Retrieved
1128:
1106:
1061:
1057:
1047:
1014:
1010:
1004:
981:Neuroscience
956:
949:
921:Bernard Katz
918:
850:
802:gap junction
783:
766:
761:
758:
729:
709:
683:
677:
647:
637:
598:
579:
557:
553:
549:
546:Bernard Katz
543:
509:
490:
487:depolarizing
482:
477:is called a
464:
448:
408:calcium ions
393:
364:
306:
289:
285:synaptic gap
284:
280:
278:
274:
266:
255:
251:active zones
239:mitochondria
224:
200:
162:
137:
95:
54:
53:
3324:Active zone
3289:Termination
3139:Interneuron
3043:Telodendron
2951:Myelination
2933:Endoneurium
2928:Perineurium
2749:Interneuron
2739:Von Economo
2687:Decussation
2682:Nerve tract
2652:Grey matter
2041:J. Neurosci
823:hippocampus
612:. Forms of
506:Termination
440:endocytosis
154:short scale
73:within the
3433:Categories
3394:Nociceptor
3134:Multipolar
3083:Nissl body
2960:Neurilemma
2923:Epineurium
2708:Cell Types
2570:Audio help
2561:2005-06-19
2266:References
1833:10447/9980
1652:Biol. Bull
1360:2008-12-26
963:Euripidean
945:exocytosis
933:exocytosis
869:Strychnine
831:amphibians
790:conductive
774:astrocytes
491:inhibitory
483:excitatory
451:taste buds
420:stochastic
400:exocytosis
173:electrical
113:exocytosis
83:perception
3409:Hair cell
2943:Neuroglia
2905:Funiculus
2794:Microglia
2767:Astrocyte
2724:Pyramidal
2677:Lemniscus
2129:CiteSeerX
1603:1945-0877
1538:1471-0080
1403:125516633
1011:Neurology
925:Paul Fatt
901:serotonin
885:endorphin
865:paralysis
819:neocortex
751:in 1963.
724:threshold
662:clonidine
365:activated
349:chemical.
258:receptors
208:dendrites
183:Structure
133:neuroglia
125:re-uptake
101:molecules
3194:Ia or Aα
3124:Unipolar
3073:Dendrite
3058:Axolemma
3053:Axoplasm
2837:Ganglion
2777:Tanycyte
2729:Purkinje
2716:Neuronal
2699:Meninges
2694:Neuropil
2572: ·
2465:15033583
2275:(2007).
2208:17068096
2151:15572174
2071:12486148
2022:15217338
1960:30728927
1952:26683686
1910:19865171
1842:17433836
1790:20495134
1688:34154835
1680:16801496
1621:29739879
1554:12384262
1546:12094216
1129:Synapses
1098:18779570
1039:38482114
1031:15985565
986:Neurexin
970:See also
909:dopamine
881:Morphine
616:include
604:include
529:Reuptake
444:recycled
410:through
404:vesicles
343:vesicles
303:Overview
71:circuits
3336:Autapse
3297:Synapse
3144:Renshaw
3119:Bipolar
2996:Neurons
2849:Ventral
2820:General
2734:Granule
2559: (
2530:minutes
2417:6261850
2408:1327434
2387:Bibcode
2366:6954549
2334:Bibcode
2256:4320287
2247:1819734
2199:2151334
2159:9527518
2106:4167209
2062:6758447
1901:2771344
1880:Bibcode
1850:1323780
1782:9651506
1672:4134559
1612:5966022
1503:6261850
1494:1327434
1473:Bibcode
1344:Cocaine
1089:2567215
1066:Bibcode
959:Verrall
905:Cocaine
889:alcohol
873:glycine
794:neurons
514:and/or
475:voltage
370:Due to
292:autapse
264:(PSD).
169:synapse
87:thought
63:muscles
59:neurons
21:synapse
3189:fibers
2827:Dorsal
2505:
2484:
2463:
2432:
2415:
2405:
2364:
2357:346205
2354:
2309:
2285:
2254:
2244:
2206:
2196:
2157:
2149:
2131:
2104:
2069:
2059:
2020:
1958:
1950:
1908:
1898:
1872:Nature
1848:
1840:
1788:
1780:
1743:
1712:
1686:
1678:
1670:
1619:
1609:
1601:
1552:
1544:
1536:
1501:
1491:
1401:
1391:
1351:
1318:
1281:
1251:
1224:
1197:
1170:
1145:9 June
1136:
1096:
1086:
1037:
1029:
961:, the
857:curare
817:, the
813:, the
811:retina
436:SNAREs
353:cells.
67:glands
44:neuron
3112:Types
3009:Parts
2878:White
2859:Ramus
2842:Ramus
2759:Glial
2155:S2CID
1956:S2CID
1928:(PDF)
1846:S2CID
1808:(PDF)
1786:S2CID
1684:S2CID
1668:JSTOR
1550:S2CID
1448:–118.
1399:S2CID
1035:S2CID
997:Notes
798:cells
516:IPSPs
512:EPSPs
204:axons
146:brain
143:human
140:adult
42:of a
3035:Axon
3017:Soma
2873:Gray
2854:Root
2832:Root
2503:ISBN
2482:ISBN
2461:PMID
2449:1662
2430:ISBN
2413:PMID
2362:PMID
2307:ISBN
2283:ISBN
2252:PMID
2204:PMID
2147:PMID
2102:PMID
2067:PMID
2018:PMID
1948:PMID
1906:PMID
1838:PMID
1778:PMID
1741:ISBN
1710:ISBN
1676:PMID
1617:PMID
1599:ISSN
1542:PMID
1534:ISSN
1499:PMID
1389:ISBN
1349:ISBN
1316:ISBN
1279:ISBN
1249:ISBN
1222:ISBN
1195:ISBN
1168:ISBN
1147:2020
1134:ISBN
1094:PMID
1027:PMID
923:and
893:GABA
829:and
827:fish
732:GABA
620:and
442:and
279:The
241:and
175:and
138:The
85:and
40:axon
3249:SVE
3244:GVE
3239:GSE
3184:SVA
3179:SSA
3174:GVA
3169:GSA
2812:PNS
2637:CNS
2453:doi
2403:PMC
2395:doi
2352:PMC
2342:doi
2242:PMC
2234:doi
2194:PMC
2186:doi
2182:578
2139:doi
2094:doi
2057:PMC
2049:doi
2010:doi
1940:doi
1936:222
1896:PMC
1888:doi
1876:461
1828:hdl
1820:doi
1770:doi
1660:doi
1656:210
1607:PMC
1591:doi
1526:doi
1489:PMC
1481:doi
1446:113
1381:doi
1312:175
1084:PMC
1074:doi
1062:105
1019:doi
897:LSD
784:An
600:of
455:ATP
290:An
206:to
127:by
65:or
3435::
2953::
2459:.
2447:.
2411:.
2401:.
2393:.
2383:33
2381:.
2377:.
2360:.
2350:.
2340:.
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2328:.
2324:.
2250:.
2240:.
2228:.
2224:.
2202:.
2192:.
2180:.
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2153:.
2145:.
2137:.
2125:47
2123:.
2100:.
2090:30
2088:.
2065:.
2055:.
2045:22
2043:.
2039:.
2016:.
2006:27
2004:.
2000:.
1954:.
1946:.
1934:.
1930:.
1904:.
1894:.
1886:.
1874:.
1870:.
1858:^
1844:.
1836:.
1826:.
1816:55
1814:.
1810:.
1784:.
1776:.
1766:26
1764:.
1733:.
1696:^
1682:.
1674:.
1666:.
1654:.
1650:.
1629:^
1615:.
1605:.
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1479:.
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1467:.
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1333:^
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1314:.
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1082:.
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1060:.
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1015:64
1013:.
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806:nm
696:.
572:.
550:pr
272:.
253:.
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1852:.
1830::
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1230:.
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27:.
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