Dendritic spine - Knowledge

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activates Cdc42, after which no feedback signaling occurs upstream to calcium and CaMKII. If tagged with monomeric-enhanced green fluorescent protein, one can see that the activation of Cdc42 is limited to just the stimulated spine of a dendrite. This is because the molecule is continuously activated during plasticity and immediately inactivates after diffusing out of the spine. Despite its compartmentalized activity, Cdc42 is still mobile out of the stimulated spine, just like RhoA. Cdc42 activates PAK, which is a protein kinase that specifically phosphorylates and, therefore, inactivates ADF/cofilin. Inactivation of cofilin leads to increased actin polymerization and expansion of the spine's volume. Activation of Cdc42 is required for this increase in spinal volume to be sustained.

412: 385: 743:, dendrites rapidly sprout and retract filopodia, small membrane organelle-lacking membranous protrusions. Recently, I-BAR protein MIM was found to contribute to the initiation process. During the first week of birth, the brain is predominated by filopodia, which eventually develop synapses. However, after this first week, filopodia are replaced by spiny dendrites but also small, stubby spines that protrude from spiny dendrites. In the development of certain filopodia into spines, filopodia recruit presynaptic contact to the dendrite, which encourages the production of spines to handle specialized postsynaptic contact with the presynaptic protrusions. 138:. Dendritic spines serve as a storage site for synaptic strength and help transmit electrical signals to the neuron's cell body. Most spines have a bulbous head (the spine head), and a thin neck that connects the head of the spine to the shaft of the dendrite. The dendrites of a single neuron can contain hundreds to thousands of spines. In addition to spines providing an anatomical substrate for memory storage and synaptic transmission, they may also serve to increase the number of possible contacts between neurons. It has also been suggested that changes in the activity of neurons have a positive effect on spine morphology. 421:

this growth, suggesting that the Rho-Rock pathway is necessary for more persistent increases in spinal volume. In addition, administration of the Cdc42 binding domain of Wasp or inhibitor targeting Pak1 activation-3 (IPA3) decreases this sustained growth in volume, demonstrating that the Cdc42-Pak pathway is needed for this growth in spinal volume as well. This is important because sustained changes in structural plasticity may provide a mechanism for the encoding, maintenance, and retrieval of memories. The observations made may suggest that Rho GTPases are necessary for these processes.

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process that stimulates actin polymerization, which in turn increases the size and shape of the spine. Large spines are more stable than smaller ones and may be resistant to modification by additional synaptic activity. Because changes in the shape and size of dendritic spines are correlated with the strength of excitatory synaptic connections and heavily depend on remodeling of its underlying actin cytoskeleton, the specific mechanisms of actin regulation, and therefore the Rho family of GTPases, are integral to the formation, maturation, and

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admitted that spines were formed during embryonic development and then would remain stable after birth. In this paradigm, variations of synaptic weight were considered as sufficient to explain memory processes at the cellular level. But since about a decade ago, new techniques of confocal microscopy demonstrated that dendritic spines are indeed motile and dynamic structures that undergo a constant turnover, even after birth.

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dendritic spine densities or sizes can affect neuronal network properties, which could lead to cognitive or mood alterations, impaired learning and memory, as well as pain hypersensitivity. Moreover, the findings suggest that maintaining spine health through therapies such as exercise, cognitive stimulation and lifestyle modifications may be helpful in preserving neuronal plasticity and improving neurological symptoms.

782:, may be resultant from abnormalities in dendritic spines, especially the number of spines and their maturity. The ratio of matured to immature spines is important in their signaling, as immature spines have impaired synaptic signaling. Fragile X syndrome is characterized by an overabundance of immature spines that have multiple filopodia in cortical dendrites. 389:

primarily by these members of the Rho family of GTPases. In the transient stage, the signaling cascade caused by synaptic activity results in LIMK1 phosphorylating ADF/cofilin via both the RhoA and Cdc42 pathways, which in turn inhibits the depolymerization of F-actin and increases the volume of the dendritic spine drastically while also inducing LTP.

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of a new skill involves a rewiring process of neural circuits. Since the extent of spine remodeling correlates with success of learning, this suggests a crucial role of synaptic structural plasticity in memory formation. In addition, changes in spine stability and strengthening occur rapidly and have been observed within hours after training.

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plasticity. On the one hand, experience and activity may drive the discrete formation of relevant synaptic connections that store meaningful information in order to allow for learning. On the other hand, synaptic connections may be formed in excess, and experience and activity may lead to the pruning of extraneous synaptic connections.

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approximation and instead uses a passive dendrite coupled to excitable spines at discrete points. Membrane dynamics in the spines are modelled using integrate and fire processes. The spike events are modelled in a discrete fashion with the wave form conventionally represented as a rectangular function.

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Overall, the evidence suggests that dendritic spines are crucial for normal brain and spinal cord function. Alterations in spine morphology may not only influence synaptic plasticity and information processing but also have a key role in many neurological diseases. Furthermore, even subtle changes in

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In lab animals of all ages, environmental enrichment has been related to dendritic branching, spine density, and overall number of synapses. In addition, skill training has been shown to lead to the formation and stabilization of new spines while destabilizing old spines, suggesting that the learning

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After the transient changes described above take place, the spine's volume decreases until it is elevated by 70 to 80 percent of the original volume. This sustained change in structural plasticity will last about thirty minutes. Once again, administration of C3 transferase and Glycyl-H1152 suppressed

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is mediated in part by the growth of new dendritic spines (or the enlargement of pre-existing spines) to reinforce a particular neural pathway. Because dendritic spines are plastic structures whose lifespan is influenced by input activity, spine dynamics may play an important role in the maintenance

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during structural plasticity brought on by long-term potentiation stimuli. Concurrent RhoA and Cdc42 activation led to a transient increase in spine growth of up to 300% for five minutes, which decayed into a smaller but sustained growth for thirty minutes. The activation of RhoA diffused around the

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Spines are particularly advantageous to neurons by compartmentalizing biochemical signals. This can help to encode changes in the state of an individual synapse without necessarily affecting the state of other synapses of the same neuron. The length and width of the spine neck has a large effect on

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In contrast, the sustained stage is focused more on activating the RhoA pathway, which ultimately results in a higher concentration of profilin, which prevents additional polymerization of actin and decreases the size of the dendritic spine from the transient stage, though still allows it to remain

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Applying a low-frequency train of two-photon glutamate uncaging in a single dendritic spine can elicit rapid activation of both RhoA and Cdc42. During the next two minutes, the volume of the stimulated spine can expand to 300 percent of its original size. However, this change in spine morphology is

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provides the theoretical framework behind the most "simple" method for modelling the flow of electrical currents along passive neural fibres. Each spine can be treated as two compartments, one representing the neck, the other representing the spine head. The compartment representing the spine head

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Theoreticians have for decades hypothesized about the potential electrical function of spines, yet our inability to examine their electrical properties has until recently stopped theoretical work from progressing too far. Recent advances in imaging techniques along with increased use of two-photon

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and spine motility has important implications for memory. If the dendritic spine is the basic unit of information storage, then the spine's ability to extend and retract spontaneously must be constrained. If not, information may be lost. Rho family of GTPases makes significant contributions to the

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Glutamate receptors (GluRs) are localized to the postsynaptic density, and are anchored by cytoskeletal elements to the membrane. They are positioned directly above their signalling machinery, which is typically tethered to the underside of the plasma membrane, allowing signals transmitted by the

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Cdc42 has been implicated in many different functions including dendritic growth, branching, and branch stability. Calcium influx into the cell through NMDA receptors binds to calmodulin and activates the Ca2+/calmodulin-dependent protein kinases II (CaMKII). In turn, CaMKII is activated and this

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on cerebellar neurons. Ramón y Cajal then proposed that dendritic spines could serve as contacting sites between neurons. This was demonstrated more than 50 years later thanks to the emergence of electron microscopy. Until the development of confocal microscopy on living tissues, it was commonly

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The SDS model was intended as a computationally simple version of the full Baer and Rinzel model. It was designed to be analytically tractable and have as few free parameters as possible while retaining those of greatest significance, such as spine neck resistance. The model drops the continuum

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Age-dependent changes in the rate of spine turnover suggest that spine stability impacts developmental learning. In youth, dendritic spine turnover is relatively high and produces a net loss of spines. This high rate of spine turnover may characterize critical periods of development and reflect

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only temporary; the volume of the spine decreases after five minutes. Administration of C3 transferase, a Rho inhibitor, or glycyl-H1152, a Rock inhibitor, inhibits the transient expansion of the spine, indicating that activation of the Rho-Rock pathway is required in some way for this process.

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lifelong learning. In addition, the formation of new spines may not significantly contribute to the connectivity of the brain, and spine formation may not bear as much of an influence on memory retention as other properties of structural plasticity, such as the increase in size of spine heads.

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There is also some evidence for loss of dendritic spines as a consequence of aging. One study using mice has noted a correlation between age-related reductions in spine densities in the hippocampus and age-dependent declines in hippocampal learning and memory. Emerging evidence has also shown

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Calcium influx through NMDA receptors activates CAMKII. CAMKII then regulates several other signaling cascades that modulate the activity of the actin-binding proteins cofilin and profilin. These cascades can be divided into two primary pathways, the RhoA and Cdc42 pathways, which are mediated

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To facilitate the analysis of interactions between many spines, Baer & Rinzel formulated a new cable theory for which the distribution of spines is treated as a continuum. In this representation, spine head voltage is the local spatial average of membrane potential in adjacent spines. The

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Despite experimental findings that suggest a role for dendritic spine dynamics in mediating learning and memory, the degree of structural plasticity's importance remains debatable. For instance, studies estimate that only a small portion of spines formed during training actually contribute to

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Experience-induced changes in dendritic spine stability also point to spine turnover as a mechanism involved in the maintenance of long-term memories, though it is unclear how sensory experience affects neural circuitry. Two general models might describe the impact of experience on structural

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Dendritic spines are small with spine head volumes ranging 0.01 μm to 0.8 μm. Spines with strong synaptic contacts typically have a large spine head, which connects to the dendrite via a membranous neck. The most notable classes of spine shape are "thin", "stubby", "mushroom", and

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leads to an increase in the rate of spine elimination and therefore impacts long-term neural circuitry. Upon restoring sensory experience after deprivation in adolescence, spine elimination is accelerated, suggesting that experience plays an important role in the net loss of spines during

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for specific brain regions. In adulthood, however, most spines remain persistent, and the half-life of spines increases. This stabilization occurs due to a developmentally regulated slow-down of spine elimination, a process which may underlie the stabilization of memories in maturity.

719:, which have a high permeability for calcium, only conduct ions if the membrane potential is sufficiently depolarized. The amount of calcium entering a spine during synaptic activity therefore depends on the depolarization of the spine head. Evidence from calcium imaging experiments ( 234:; without a dynamic cytoskeleton, spines would be unable to rapidly change their volumes or shapes in responses to stimuli. These changes in shape might affect the electrical properties of the spine. The cytoskeleton of dendritic spines is primarily made of filamentous actin ( 462:

of its synapsing axon and comprises ~10% of the spine's membrane surface area; neurotransmitters released from the active zone bind receptors in the postsynaptic density of the spine. Half of the synapsing axons and dendritic spines are physically tethered by

547:. Furthermore, spine number is very variable and spines come and go; in a matter of hours, 10-20% of spines can spontaneously appear or disappear on the pyramidal cells of the cerebral cortex, although the larger "mushroom"-shaped spines are the most stable. 656:. Because significant changes in spine density occur in various brain and spinal cord diseases, this suggests a balanced state of spine dynamics in normal circumstances, which may be susceptible to disequilibrium under varying pathological conditions. 746:

Spines, however, require maturation after formation. Immature spines have impaired signaling capabilities, and typically lack "heads" (or have very small heads), only necks, while matured spines maintain both heads and necks.

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are present. Because spines have a cytoskeleton of primarily actin, this allows them to be highly dynamic in shape and size. The actin cytoskeleton directly determines the morphology of the spine, and actin regulators, small

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Kiss C, Li J, Szeles A, Gizatullin RZ, Kashuba VI, Lushnikova T, et al. (1 January 1997). "Assignment of the ARHA and GPX1 genes to human chromosome bands 3p21.3 by in situ hybridization and with somatic cell hybrids".

636:, a marked increase in structural plasticity occurs near the trauma site, and a five- to eightfold increase from control rates in spine turnover has been observed. Dendrites disintegrate and reassemble rapidly during 483:. The localization of signalling elements to their GluRs is particularly important in ensuring signal cascade activation, as GluRs would be unable to affect particular downstream effects without nearby signallers. 393:

Murakoshi, Wang, and Yasuda (2011) examined the effects of Rho GTPase activation on the structural plasticity of single dendritic spines elucidating differences between the transient and sustained phases.

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studies have shown that there is a continuum of shapes between these categories. The variable spine shape and volume is thought to be correlated with the strength and maturity of each spine-synapse.

335:, a protein that also modulates the regulation and timing of cell division. In the context of activity in neurons, RhoA is activated in the following manner: once calcium has entered a cell through 358:

A study conducted by Murakoshi et al. in 2011 implicated the Rho GTPases RhoA and Cdc42 in dendritic spine morphogenesis. Both GTPases were quickly activated in single dendritic spines of

355:. Cofilin's function is to reorganize the actin cytoskeleton of a cell; namely, it depolymerizes actin segments and thus inhibits the growth of growth cones and the repair of axons. 29: 698:

formulation maintains the feature that there is no direct electrical coupling between neighboring spines; voltage spread along dendrites is the only way for spines to interact.

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vicinity of the spine undergoing stimulation, and it was determined that RhoA is necessary for the transient phase and most likely the sustained phase as well of spine growth.

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Ngo-Anh TJ, Bloodgood BL, Lin M, Sabatini BL, Maylie J, Adelman JP (May 2005). "SK channels and NMDA receptors form a Ca2+-mediated feedback loop in dendritic spines".

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glutamate uncaging have led to a wealth of new discoveries; we now suspect that there are voltage-dependent sodium, potassium, and calcium channels in the spine heads.

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dendritic spine abnormalities in the pain processing regions of the spinal cord nociceptive system, including superficial and intermediate zones of the dorsal horn.

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is also expressed on the spine surface, and is believed to play a role in spine survival. The tip of the spine contains an electron-dense region referred to as the "

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von Bohlen und Halbach O, Zacher C, Gass P, Unsicker K (March 2006). "Age-related alterations in hippocampal spines and deficiencies in spatial memory in mice".

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Signalling from GluRs is mediated by the presence of an abundance of proteins, especially kinases, that are localized to the postsynaptic density. These include

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Dendritic spines are very "plastic", that is, spines change significantly in shape, volume, and number in small time courses. Because spines have a primarily

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Tashiro A, Yuste R (July 2004). "Regulation of dendritic spine motility and stability by Rac1 and Rho kinase: evidence for two forms of spine motility".

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learning capacity in adolescence—different cortical areas exhibit differing levels of synaptic turnover during development, possibly reflecting varying

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Brown CE, Murphy TH (April 2008). "Livin' on the edge: imaging dendritic spine turnover in the peri-infarct zone during ischemic stroke and recovery".

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In addition to their electrophysiological activity and their receptor-mediated activity, spines appear to be vesicularly active and may even translate

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Holtmaat A, Wilbrecht L, Knott GW, Welker E, Svoboda K (June 2006). "Experience-dependent and cell-type-specific spine growth in the neocortex".

347:, which leads to the activation of RhoA. The activation of the RhoA protein will activate ROCK, a RhoA kinase, which leads to the stimulation of 629:

development. In addition, other sensory deprivation paradigms—such as whisker trimming—have been shown to increase the stability of new spines.

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Benson, Curtis A.; Fenrich, Keith K.; Olson, Kai-Lan; Patwa, Siraj; Bangalore, Lakshmi; Waxman, Stephen G.; Tan, Andrew M. (2020-05-27).

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Kasai H, Matsuzaki M, Noguchi J, Yasumatsu N, Nakahara H (July 2003). "Structure-stability-function relationships of dendritic spines".

1585:"Dendritic BDNF synthesis is required for late-phase spine maturation and recovery of cortical responses following sensory deprivation" 314:, either in globular (G-actin) or filamentous (F-actin) forms. The role of Rho family of GTPases and its effects in the stability of 3596: 566:

and confocal microscopy have shown that spine volume changes depending on the types of stimuli that are presented to a synapse.

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onto their equally numerous spines, whereas the number of spines on Purkinje neuron dendrites is an order of magnitude larger.

167:. Excitatory axon proximity to dendritic spines is not sufficient to predict the presence of a synapse, as demonstrated by the 1758:"Deafening drives cell-type-specific changes to dendritic spines in a sensorimotor nucleus important to learned vocalizations" 3340: 3321: 2413:"Dendritic spine remodeling following early and late Rac1 inhibition after spinal cord injury: evidence for a pain biomarker" 3408:

Lynch G, Rex CS, Gall CM (January 2007). "LTP consolidation: substrates, explanatory power, and functional significance".

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Zhao, Peng; Hill, Myriam; Liu, Shujun; Chen, Lubin; Bangalore, Lakshmi; Waxman, Stephen G.; Tan, Andrew M. (2016-06-01).

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Benson, Curtis A.; King, Jared F.; Reimer, Marike L.; Kauer, Sierra D.; Waxman, Stephen G.; Tan, Andrew M. (2022-12-03).

543:, they are dynamic, and the majority of spines change their shape within seconds to minutes because of the dynamicity of 3525:

Yuste R, Majewska A, Holthoff K (July 2000). "From form to function: calcium compartmentalization in dendritic spines".

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Research in neurological diseases and injuries shed further light on the nature and importance of spine turnover. After

411: 384: 640:—as with stroke, survivors showed an increase in dendritic spine turnover. While a net loss of spines is observed in 2761:

Baer SM, Rinzel J (April 1991). "Propagation of dendritic spikes mediated by excitable spines: a continuum theory".

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Tackenberg C, Ghori A, Brandt R (June 2009). "Thin, stubby or mushroom: spine pathology in Alzheimer's disease".

2722:"Local postsynaptic voltage-gated sodium channel activation in dendritic spines of olfactory bulb granule cells" 727:

indicates that spines with high resistance necks experience larger calcium transients during synaptic activity.

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Conversely, while enrichment and training are related to increases in spine formation and stability, long-term

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Calabrese B, Wilson MS, Halpain S (February 2006). "Development and regulation of dendritic spine synapses".

1158:"Dynamic actin filaments are required for stable long-term potentiation (LTP) in area CA1 of the hippocampus" 1056:

Kapitein LC, Schlager MA, Kuijpers M, Wulf PS, van Spronsen M, MacKintosh FC, Hoogenraad CC (February 2010).

648:, cocaine and amphetamine use have been linked to increases in dendritic branching and spine density in the 294:

have also been identified in spines, supporting the vesicular activity in dendritic spines. The presence of

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Gray EG (June 1959). "Electron microscopy of synaptic contacts on dendrite spines of the cerebral cortex".

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in spines also suggests protein translational activity in the spine itself, not just in the dendrite.

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Holtmaat AJ, Trachtenberg JT, Wilbrecht L, Shepherd GM, Zhang X, Knott GW, Svoboda K (January 2005).

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Bressloff PC, Coombes S (2000). "Solitary Waves in a Model of Dendritic Cable with Active Spines".

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Yuste R, Denk W (June 1995). "Dendritic spines as basic functional units of neuronal integration".

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the degree of compartmentalization, with thin spines being the most biochemically isolated spines.

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Bywalez WG, Patirniche D, Rupprecht V, Stemmler M, Herz AV, Pálfi D, et al. (February 2015).

956:"Preventing adolescent synaptic pruning in mouse prelimbic cortex via local knockdown of α4βδ GABA 4415: 3854: 3582: 775: 724: 645: 163:

from axons, although sometimes both inhibitory and excitatory connections are made onto the same

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896:"Ultrastructural analysis of dendritic spine necks reveals a continuum of spine morphologies" 720: 507: 3496:

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Spine maintenance and plasticity is activity-dependent and activity-independent.

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to be further propagated by their nearby signalling elements to activate

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Dendritic spines were first described at the end of the 19th century by

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Calcium transients in spines are a key trigger for synaptic plasticity.

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and is believed to play an important role in calcium handling. "Smooth"

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The cytoskeleton of dendritic spines is particularly important in their

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The morphogenesis of dendritic spines is critical to the induction of

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may receive tens of thousands of mostly excitatory inputs from other

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Small protrusion on a dendrite that receives input from a single axon

3102: 1916:"Transient and persistent dendritic spines in the neocortex in vivo" 735:

Dendritic spines can develop directly from dendritic shafts or from

282:(SERs) have been identified in dendritic spines. Formation of this " 4049: 4034: 4029: 3753: 3748: 3675: 3670: 2636: 637: 592: 531: 471:, which forms cell-to-cell adherent junctions between two neurons. 468: 195: 175: 127: 34: 28: 3538: 2995:"MIM-Induced Membrane Bending Promotes Dendritic Spine Initiation" 2719: 1542:

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Grunditz A, Holbro N, Tian L, Zuo Y, Oertner TG (December 2008).

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Ofer N, Berger DR, Kasthuri N, Lichtman JW, Yuste R (July 2021).

755:

Emerging research has indicate abnormalities in spine density in

579: 487: 476: 464: 352: 275: 239: 235: 219: 135: 2992: 331:

One of the major Rho GTPases involved in spine morphogenesis is

3972: 3927: 3692: 771: 633: 596: 252: 179: 3445:

Matsuzaki M, Honkura N, Ellis-Davies GC, Kasai H (June 2004).

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Bhatt DH, Zhang S, Gan WB (2009). "Dendritic spine dynamics".

1325: 2076:

Brown CE, Li P, Boyd JD, Delaney KR, Murphy TH (April 2007).

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Roberts TF, Tschida KA, Klein ME, Mooney R (February 2010).

1375:"Actin in dendritic spines: connecting dynamics to function" 1055: 379: 310:(LTP). The morphology of the spine depends on the states of 207: 4011: 3084: 2851:"Spine Ca2+ signaling in spike-timing-dependent plasticity" 2622: 763: 551: 451: 447: 332: 268: 260: 131: 893: 206:. Dendritic spines occur at a density of up to 5 spines/1 3495: 3257:"Spine motility. Phenomenology, mechanisms, and function" 3087:"Dendritic spine pathology in neuropsychiatric disorders" 2564:

Araya R, Nikolenko V, Eisenthal KB, Yuste R (July 2007).

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at an elevated level compared to an unpotentiated spine.

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Kaneko M, Xie Y, An JJ, Stryker MP, Xu B (April 2012).

2353: 1506: 1421: 3524: 3316:(Third ed.). New York: Oxford University Press. 2949: 2897: 2210: 1155: 858: 554:

partially determines spine levels, and low levels of

3027: 1368: 710: 692: 583:

Experience-dependent spine formation and elimination

569: 3034:Pelucchi S, Stringhi R, Marcello E (January 2020). 1804: 1268: 1264: 1262: 1260: 1258: 1256: 1254: 1017:"Saturated Reconstruction of a Volume of Neocortex" 1008: 126:) is a small membranous protrusion from a neuron's 3330: 3309: 2514: 2467: 2075: 1582: 960:receptors increases anxiety response in adulthood" 954:Evrard MR, Li M, Shen H, Smith SS (October 2021). 271:results in consistently smaller dendritic spines. 3335:. Baltimore: The Johns Hopkins University Press. 2945: 2943: 1535: 953: 4407: 3307: 2795: 1805:De Roo M, Klauser P, Muller D (September 2008). 1251: 323:of dendritic spines and to learning and memory. 3370: 3254: 2410: 1636: 823: 267:, rapidly modify this cytoskeleton. Overactive 2940: 1864: 1457: 1156:Krucker T, Siggins GR, Halpain S (June 2000). 225: 3590: 3133: 2848: 2515:Harris KM, Fiala JC, Ostroff L (April 2003). 2170: 1755: 3407: 2986: 1865:Zuo Y, Lin A, Chang P, Gan WB (April 2005). 1269:Murakoshi H, Wang H, Yasuda R (April 2011). 1214: 701: 130:that typically receives input from a single 3203: 3040:International Journal of Molecular Sciences 2124: 1049: 852: 3597: 3583: 2760: 2715: 2713: 2566:"Sodium channels amplify spine potentials" 1967: 1694: 1692: 1690: 1576: 689:alone should carry the active properties. 498:(calmodulin-dependent protein kinase II), 407:Sustained changes in structural plasticity 398:Transient changes in structural plasticity 3478: 3272: 3231: 3221: 3110: 3061: 3051: 3010: 2923: 2874: 2809: 2737: 2665: 2599: 2589: 2540: 2444: 2387: 2330: 2101: 2001: 1931: 1882: 1832: 1822: 1781: 1732: 1670: 1608: 1559: 1483: 1458:Scott EK, Reuter JE, Luo L (April 2003). 1398: 1302: 1191: 1181: 1132: 1122: 1104: 1073: 1032: 991: 919: 819: 817: 815: 813: 811: 809: 807: 574: 380:Observed changes in structural plasticity 3510:10.1146/annurev.physiol.64.081501.160008 3331:Sudhof TC, Stevens CF, Cowan WM (2001). 3255:Bonhoeffer T, Yuste R (September 2002). 2166: 2164: 1907: 750: 578: 410: 383: 3570:Spiny Dendrite - Cell Centered Database 2710: 1968:Yang G, Pan F, Gan WB (December 2009). 1963: 1961: 1959: 1749: 1687: 667: 4408: 3312:The Neuron: Cell and Molecular Biology 1860: 1858: 1856: 1854: 1852: 838:10.1146/annurev.neuro.30.051606.094222 804: 562:encourages spine growth. Furthermore, 458:" (PSD). The PSD directly apposes the 3578: 2206: 2204: 2202: 2185:10.1146/annurev.physiol.010908.163140 2161: 1632: 1630: 1628: 351:, which in turn inhibits the protein 3152: 2849:Nevian T, Sakmann B (October 2006). 1956: 1217:Molecular and Cellular Neurosciences 949: 947: 564:two-photon laser scanning microscopy 429: 2798:SIAM Journal on Applied Mathematics 1849: 1756:Tschida KA, Mooney R (March 2012). 13: 3604: 3300: 3223:10.1523/JNEUROSCI.16-09-02983.1996 2213:"Dendritic Spines and Pain Memory" 2199: 1625: 1476:10.1523/JNEUROSCI.23-08-03118.2003 1124:10.1523/JNEUROSCI.19-11-04314.1999 587:Spine plasticity is implicated in 246:(MAPs) are present, and organized 14: 4432: 3563: 3308:Levitan IB, Kaczmarek LK (2002). 944: 711:Modeling spine calcium transients 693:Baer and Rinzel's continuum model 570:Importance to learning and memory 159:Dendritic spines usually receive 49:Common types of dendritic spines. 3422:10.1016/j.neuropharm.2006.07.027 3204:Dailey ME, Smith SJ (May 1996). 2256:Journal of Neuroscience Research 370: 301: 102:Anatomical terms of microanatomy 43: 27: 3761:Oligodendrocyte progenitor cell 3248: 3197: 3146: 3127: 3078: 2952:Current Opinion in Neurobiology 2891: 2842: 2789: 2754: 2659: 2616: 2557: 2508: 2461: 2404: 2347: 2290: 2247: 2118: 2069: 2018: 1798: 1500: 1451: 1415: 1362: 1319: 1105:Kim CH, Lisman JE (June 1999). 326: 244:microtubule-associated proteins 154: 2916:10.1523/JNEUROSCI.2702-08.2008 2867:10.1523/JNEUROSCI.1749-06.2006 2315:10.1523/JNEUROSCI.2858-19.2020 2094:10.1523/JNEUROSCI.4295-06.2007 1601:10.1523/JNEUROSCI.4462-11.2012 1425:Cytogenetics and Cell Genetics 1208: 1149: 1098: 887: 730: 1: 3385:10.1016/S0166-4328(02)00271-1 3274:10.1016/s0896-6273(02)00906-6 1340:10.1016/S0166-2236(03)00162-0 826:Annual Review of Neuroscience 798: 521: 424: 362:in the CA1 region of the rat 3136:Rev. Trim. Histol. Norm. Pat 3012:10.1016/j.devcel.2015.04.014 2739:10.1016/j.neuron.2014.12.051 1933:10.1016/j.neuron.2005.01.003 1884:10.1016/j.neuron.2005.04.001 1824:10.1371/journal.pbio.0060219 1774:10.1016/j.neuron.2011.12.038 762:Cognitive disorders such as 481:signal transduction cascades 280:smooth endoplasmic reticulum 182:in the brain, including the 141: 7: 3871:Postganglionic nerve fibers 3498:Annual Review of Physiology 3210:The Journal of Neuroscience 2904:The Journal of Neuroscience 2855:The Journal of Neuroscience 2303:The Journal of Neuroscience 2173:Annual Review of Physiology 2082:The Journal of Neuroscience 1589:The Journal of Neuroscience 1464:The Journal of Neuroscience 1379:The Journal of Cell Biology 1111:The Journal of Neuroscience 676: 604:of memory over a lifetime. 226:Cytoskeleton and organelles 10: 4437: 4421:Computational neuroscience 3866:Preganglionic nerve fibers 3373:Behavioural Brain Research 2964:10.1016/j.conb.2009.05.013 2763:Journal of Neurophysiology 2474:Journal of Neurophysiology 2417:Journal of Neurophysiology 1521:10.1152/physiol.00042.2005 1034:10.1016/j.cell.2015.06.054 984:10.1038/s41598-021-99965-8 900:Developmental Neurobiology 873:10.2174/156720509788486554 861:Current Alzheimer Research 785: 525: 4376:Olfactory receptor neuron 4331: 4272: 4265: 4201: 4131: 4088: 4048: 4040:Neurofibril/neurofilament 4010: 3992: 3985: 3971: 3918: 3890: 3796: 3787: 3734: 3691: 3684: 3621: 3612: 2820:10.1137/s0036139999356600 2225:10.1177/10738584221138251 1229:10.1016/j.mcn.2004.04.001 1075:10.1016/j.cub.2009.12.052 702:Spike-diffuse-spike model 434:Dendritic spines express 286:" depends on the protein 100: 88: 76: 64: 59: 54: 42: 26: 21: 2775:10.1152/jn.1991.65.4.874 2372:10.1177/1744806916688016 2139:10.1177/1073858407309854 446:) on their surface. The 174:Spines are found on the 2591:10.1073/pnas.0705282104 1328:Trends in Neurosciences 776:intellectual disability 725:compartmental modelling 646:intellectual disability 278:. Stacked discs of the 4323:Neuromuscular junction 4186:III or Aδ or fast pain 2533:10.1098/rstb.2002.1254 1183:10.1073/pnas.100139797 792:Santiago Ramón y Cajal 584: 575:Evidence of importance 417: 390: 308:long-term potentiation 2486:10.1152/jn.00095.2009 2429:10.1152/jn.01057.2015 1561:10.1093/cercor/bhm041 1391:10.1083/jcb.201003008 751:Clinical significance 721:two-photon microscopy 582: 508:Protein Phosphatase-1 414: 387: 210:stretch of dendrite. 4341:Meissner's corpuscle 4306:Postsynaptic density 4203:Efferent nerve fiber 4191:IV or C or slow pain 4133:Afferent nerve fiber 3959:Satellite glial cell 3053:10.3390/ijms21030908 668:Importance contested 506:(Protein Kinase A), 502:(Protein Kinase C), 456:postsynaptic density 192:medium spiny neurons 37:medium spiny neuron. 33:Spiny dendrite of a 4346:Merkel nerve ending 3527:Nature Neuroscience 3471:10.1038/nature02617 3463:2004Natur.429..761M 3167:1959Natur.183.1592G 3161:(4675): 1592–1593. 3091:Nature Neuroscience 2910:(50): 13457–13466. 2861:(43): 11001–11013. 2680:1995Natur.375..682Y 2625:Nature Neuroscience 2582:2007PNAS..10412347A 2576:(30): 12347–12352. 2366:: 174480691668801. 2219:: 107385842211382. 2047:10.1038/nature04783 2039:2006Natur.441..979H 1994:10.1038/nature08577 1986:2009Natur.462..920Y 1725:10.1038/nature08759 1717:2010Natur.463..948R 1663:10.1038/nature08389 1655:2009Natur.462..915X 1295:10.1038/nature09823 1287:2011Natur.472..100M 1174:2000PNAS...97.6856K 976:2021NatSR..1121059E 768:Alzheimer's disease 737:dendritic filopodia 642:Alzheimer's disease 626:sensory deprivation 528:Synaptic plasticity 512:Fyn tyrosine kinase 436:glutamate receptors 232:synaptic plasticity 149:Electron microscopy 4381:Photoreceptor cell 4351:Pacinian corpuscle 4282:Electrical synapse 4236:Lower motor neuron 4231:Upper motor neuron 3952:Internodal segment 3892:Connective tissues 3862:Autonomic ganglion 2999:Developmental Cell 2217:The Neuroscientist 2127:The Neuroscientist 964:Scientific Reports 912:10.1002/dneu.22829 780:fragile X syndrome 585: 418: 391: 178:of most principal 95:H2.00.06.1.00036 71:gemmula dendritica 4403: 4402: 4399: 4398: 4366:Free nerve ending 4333:Sensory receptors 4261: 4260: 4176:Ib or Golgi or Aα 4084: 4083: 3967: 3966: 3844:Ramus communicans 3783: 3782: 3779: 3778: 3649:Commissural fiber 3644:Association fiber 3639:Projection fibers 3457:(6993): 761–766. 3410:Neuropharmacology 3342:978-0-8018-6498-8 3323:978-0-19-514522-9 3175:10.1038/1831592a0 2674:(6533): 682–684. 2527:(1432): 745–748. 2309:(22): 4297–4308. 2268:10.1002/jnr.20759 2088:(15): 4101–4109. 2033:(7096): 979–983. 1980:(7275): 920–924. 1711:(7283): 948–952. 1649:(7275): 915–919. 1595:(14): 4790–4802. 1437:10.1159/000134729 1281:(7341): 100–104. 1168:(12): 6856–6861. 1117:(11): 4314–4324. 654:nucleus accumbens 650:prefrontal cortex 599:. In particular, 430:Receptor activity 360:pyramidal neurons 216:pyramidal neurons 184:pyramidal neurons 116: 115: 111: 4428: 4296:Synaptic vesicle 4291:Chemical synapse 4270: 4269: 3990: 3989: 3983: 3982: 3794: 3793: 3689: 3688: 3619: 3618: 3599: 3592: 3585: 3576: 3575: 3558: 3521: 3492: 3482: 3441: 3404: 3367: 3346: 3327: 3315: 3295: 3294: 3276: 3267:(6): 1019–1027. 3252: 3246: 3245: 3235: 3225: 3216:(9): 2983–2994. 3201: 3195: 3194: 3150: 3144: 3143: 3131: 3125: 3124: 3114: 3082: 3076: 3075: 3065: 3055: 3031: 3025: 3024: 3014: 2990: 2984: 2983: 2947: 2938: 2937: 2927: 2895: 2889: 2888: 2878: 2846: 2840: 2839: 2813: 2793: 2787: 2786: 2758: 2752: 2751: 2741: 2717: 2708: 2707: 2688:10.1038/375682a0 2663: 2657: 2656: 2620: 2614: 2613: 2603: 2593: 2561: 2555: 2554: 2544: 2512: 2506: 2505: 2480:(4): 2396–2409. 2465: 2459: 2458: 2448: 2423:(6): 2893–2910. 2408: 2402: 2401: 2391: 2351: 2345: 2344: 2334: 2294: 2288: 2287: 2251: 2245: 2244: 2208: 2197: 2196: 2168: 2159: 2158: 2122: 2116: 2115: 2105: 2073: 2067: 2066: 2022: 2016: 2015: 2005: 1965: 1954: 1953: 1935: 1911: 1905: 1904: 1886: 1862: 1847: 1846: 1836: 1826: 1802: 1796: 1795: 1785: 1768:(5): 1028–1039. 1753: 1747: 1746: 1736: 1696: 1685: 1684: 1674: 1634: 1623: 1622: 1612: 1580: 1574: 1573: 1563: 1539: 1533: 1532: 1504: 1498: 1497: 1487: 1470:(8): 3118–3123. 1455: 1449: 1448: 1431:(3–4): 228–230. 1419: 1413: 1412: 1402: 1366: 1360: 1359: 1323: 1317: 1316: 1306: 1266: 1249: 1248: 1212: 1206: 1205: 1195: 1185: 1153: 1147: 1146: 1136: 1126: 1102: 1096: 1095: 1077: 1053: 1047: 1046: 1036: 1012: 1006: 1005: 995: 951: 942: 941: 923: 891: 885: 884: 856: 850: 849: 821: 610:critical periods 601:long-term memory 545:actin remodeling 161:excitatory input 108:edit on Wikidata 105: 47: 31: 19: 18: 4436: 4435: 4431: 4430: 4429: 4427: 4426: 4425: 4406: 4405: 4404: 4395: 4327: 4257: 4206: 4197: 4181:II or Aβ and Aγ 4136: 4127: 4080: 4070:Apical dendrite 4065:Dendritic spine 4044: 4006: 3976: 3963: 3947:Node of Ranvier 3942:Myelin incisure 3914: 3886: 3775: 3766:Oligodendrocyte 3749:Ependymal cells 3730: 3680: 3608: 3603: 3566: 3561: 3343: 3324: 3303: 3301:Further reading 3298: 3253: 3249: 3202: 3198: 3151: 3147: 3132: 3128: 3103:10.1038/nn.2741 3083: 3079: 3032: 3028: 2991: 2987: 2948: 2941: 2896: 2892: 2847: 2843: 2811:10.1.1.104.1307 2794: 2790: 2759: 2755: 2718: 2711: 2664: 2660: 2621: 2617: 2562: 2558: 2513: 2509: 2466: 2462: 2409: 2405: 2352: 2348: 2295: 2291: 2252: 2248: 2209: 2200: 2169: 2162: 2123: 2119: 2074: 2070: 2023: 2019: 1966: 1957: 1912: 1908: 1863: 1850: 1803: 1799: 1754: 1750: 1697: 1688: 1635: 1626: 1581: 1577: 1548:Cerebral Cortex 1540: 1536: 1505: 1501: 1456: 1452: 1420: 1416: 1367: 1363: 1324: 1320: 1267: 1252: 1213: 1209: 1154: 1150: 1103: 1099: 1062:Current Biology 1054: 1050: 1013: 1009: 959: 952: 945: 892: 888: 857: 853: 822: 805: 801: 788: 753: 733: 713: 704: 695: 679: 670: 577: 572: 534: 524: 475:GluRs into the 432: 427: 409: 400: 382: 373: 329: 304: 284:spine apparatus 228: 157: 144: 120:dendritic spine 112: 50: 38: 22:Dendritic spine 17: 12: 11: 5: 4434: 4424: 4423: 4418: 4416:Neurohistology 4401: 4400: 4397: 4396: 4394: 4393: 4391:Taste receptor 4388: 4383: 4378: 4373: 4368: 4363: 4361:Muscle spindle 4358: 4356:Ruffini ending 4353: 4348: 4343: 4337: 4335: 4329: 4328: 4326: 4325: 4320: 4318:Ribbon synapse 4315: 4310: 4309: 4308: 4303: 4298: 4288: 4278: 4276: 4267: 4263: 4262: 4259: 4258: 4256: 4255: 4254: 4253: 4248: 4243: 4233: 4228: 4223: 4218: 4212: 4210: 4199: 4198: 4196: 4195: 4194: 4193: 4188: 4183: 4178: 4173: 4163: 4158: 4153: 4148: 4142: 4140: 4138:Sensory neuron 4129: 4128: 4126: 4125: 4124: 4123: 4113: 4108: 4106:Pseudounipolar 4103: 4098: 4092: 4090: 4086: 4085: 4082: 4081: 4079: 4078: 4077: 4076: 4074:Basal dendrite 4067: 4062: 4054: 4052: 4046: 4045: 4043: 4042: 4037: 4032: 4027: 4025:Axon terminals 4022: 4016: 4014: 4008: 4007: 4005: 4004: 3998: 3996: 3987: 3980: 3969: 3968: 3965: 3964: 3962: 3961: 3956: 3955: 3954: 3949: 3944: 3939: 3924: 3922: 3916: 3915: 3913: 3912: 3907: 3902: 3896: 3894: 3888: 3887: 3885: 3884: 3879: 3877:Nerve fascicle 3874: 3868: 3859: 3858: 3857: 3852: 3840: 3839: 3838: 3833: 3823: 3822: 3821: 3816: 3811: 3800: 3798: 3791: 3785: 3784: 3781: 3780: 3777: 3776: 3774: 3773: 3768: 3763: 3758: 3757: 3756: 3746: 3740: 3738: 3732: 3731: 3729: 3728: 3723: 3718: 3713: 3708: 3703: 3697: 3695: 3686: 3682: 3681: 3679: 3678: 3673: 3668: 3667: 3666: 3661: 3656: 3651: 3646: 3641: 3631: 3625: 3623: 3616: 3610: 3609: 3606:Nervous tissue 3602: 3601: 3594: 3587: 3579: 3573: 3572: 3565: 3564:External links 3562: 3560: 3559: 3533:(7): 653–659. 3522: 3493: 3442: 3405: 3379:(1–2): 87–95. 3368: 3358:(5): 159–164. 3347: 3341: 3328: 3322: 3304: 3302: 3299: 3297: 3296: 3247: 3196: 3145: 3126: 3097:(3): 285–293. 3077: 3026: 3005:(6): 644–659. 2985: 2939: 2890: 2841: 2804:(2): 432–453. 2788: 2769:(4): 874–890. 2753: 2732:(3): 590–601. 2709: 2658: 2637:10.1038/nn1449 2631:(5): 642–649. 2615: 2556: 2507: 2460: 2403: 2360:Molecular Pain 2346: 2289: 2262:(4): 525–531. 2246: 2198: 2160: 2133:(2): 139–146. 2117: 2068: 2017: 1955: 1926:(2): 279–291. 1906: 1877:(2): 181–189. 1848: 1797: 1748: 1686: 1624: 1575: 1554:(1): 151–161. 1534: 1499: 1450: 1414: 1385:(4): 619–629. 1369:Hotulainen P, 1361: 1334:(7): 360–368. 1318: 1250: 1223:(3): 429–440. 1207: 1148: 1097: 1048: 1027:(3): 648–661. 1007: 957: 943: 906:(5): 746–757. 886: 851: 802: 800: 797: 787: 784: 752: 749: 741:synaptogenesis 732: 729: 717:NMDA receptors 712: 709: 703: 700: 694: 691: 678: 675: 669: 666: 576: 573: 571: 568: 560:NMDA receptors 523: 520: 431: 428: 426: 423: 408: 405: 399: 396: 381: 378: 372: 369: 343:and activates 339:, it binds to 337:NMDA receptors 328: 325: 303: 300: 227: 224: 200:Purkinje cells 156: 153: 147:"bifurcated". 143: 140: 114: 113: 104: 98: 97: 92: 86: 85: 80: 74: 73: 68: 62: 61: 57: 56: 52: 51: 48: 40: 39: 32: 24: 23: 15: 9: 6: 4: 3: 2: 4433: 4422: 4419: 4417: 4414: 4413: 4411: 4392: 4389: 4387: 4384: 4382: 4379: 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