Spatiotemporal dynamics of dendritic spines in the living brain
Open accessAbstract
Dendritic spines are ubiquitous postsynaptic sites of most excitatory synapses in the mammalian brain, and thus may serve as structural indicators of functional synapses. Recent works have suggested that neuronal coding of memories may be associated with rapid alterations in spine formation and elimination. Technological advances have enabled researchers to study spine dynamics in vivo during development as well as under various physiological and pathological conditions. We believe that better understanding of the spatiotemporal patterns of spine dynamics will help elucidate the principles of experience-dependent circuit modification and information processing in the living brain.
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Written from the paper’s full text.
TL;DR
This review synthesizes in vivo imaging studies showing that dendritic spines are constantly formed and eliminated in the living brain, with the balance between these processes shifting across development, sensory experience, learning, and disease. It argues that understanding the spatiotemporal patterns of spine dynamics is key to elucidating experience-dependent circuit modification and information storage.
Key findings
- Spine density changes non-monotonically over development: motility and turnover decrease between P7 and P24 in rodent barrel cortex while density rises, then elimination outpaces formation — between P28 and P42, 17% of spines are eliminated along apical dendrites of layer 5 pyramidal neurons in mouse barrel cortex versus only 5% formed (Zuo et al., 2005a,b). Large-headed spines are more stable than thin ones, and newly formed spines are more likely to be eliminated than pre-existing ones.
- Sensory manipulations alter spine dynamics in a paradigm- and cell-type-specific manner: unilateral whisker trimming in 1-month-old mice reduced spine elimination without perturbing formation, an effect mimicked by NMDA receptor blockade; chessboard trimming promoted turnover and selectively stabilized new spines on layer 5 pyramidal neurons with complex apical tufts; monocular deprivation increased spine formation on layer 5 but not layer 2/3 neurons in binocular cortex.
- Learning induces rapid spine formation followed by elevated elimination that returns density to baseline; the amount of spines gained during initial motor learning correlates with acquisition performance, and survival of new spines correlates with skill retention. In fear conditioning, auditory cortex showed increased formation with paired conditioning and increased elimination with unpaired conditioning, while frontal association cortex showed the opposite pattern for learning versus extinction.
- Spine changes occur at spatially selective "hot spots": new spines formed during repeated training of the same motor task cluster, whereas spines formed during different tasks or motor enrichment do not; a spine eliminated during fear conditioning is typically replaced within 2 μm during extinction. Disease models show distinct signatures — rapid reversible spine loss in ischemia (reversible if reperfused within 20–60 min), accelerated spine loss near β-amyloid plaques, and increased turnover with failure of new spine persistence in Huntington's and MECP2-duplication models.
Methods
This is a narrative review, not a primary study. It synthesizes results from in vivo time-lapse imaging studies (predominantly multi-photon microscopy, plus whole-cell patch recording and single-spine calcium imaging) across rodent cortex, songbird, and disease models, organized thematically by development, sensory experience, learning, disease, glial contribution, and spatial patterning. No new data, sample, or statistical analysis are reported.
Limitations
The paper states that all reviewed work focuses on the postsynaptic side only, and that identifying the presynaptic partner of an imaged spine remains a technical challenge; it also notes that declarative memory is difficult to study because the hippocampus is beyond the reach of standard two-photon microscopy and declarative memories are diffusely stored. Evident from the text: as a review, it presents no primary data or quantitative synthesis, and several cited findings (e.g., clustering rules, glial roles) rest on single studies whose replication status is not discussed.
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Claims the paper makes
0 of 124 cited sources checked · 0 supported · 0 flagged
- Premisecitation
Stable spines mostly represent synaptic contacts.
“While stable spines mostly represent synaptic contacts, only a small fraction of transient spines represent short-lived synaptic contacts, and the rest of them represent failed synaptogenesis (Trachtenberg et al., 2002; Knott et al., 2006; Cane et al., 2014).”
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- cites Knott et al., 2006 source
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- cites Cane et al., 2014 source
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- The authors state this as an assumption: In principle, spine dynamics represent synapse dynamics.
- Premisecitation
In 2002, two laboratories demonstrated the possibility to track the same spine in the living brain over weeks.
“In 2002, the pioneering work from two laboratories (Grutzendler et al., 2002; Trachtenberg et al., 2002) demonstrated the possibility to track the same spine in the living brain over a long period (i.e., weeks) of time.”
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Spines and their structural dynamics may serve as indicators for synaptic connectivity and modifications thereof.
“Therefore, spines and their structural dynamics may serve as indicators for synaptic connectivity and modifications thereof (Segal, 2005; Tada and Sheng, 2006; Harms and Dunaevsky, 2007).”
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- cites Tada and Sheng, 2006
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- cites Harms and Dunaevsky, 2007
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- Rests on an assumption: Dendritic spines are the postsynaptic sites of the great majority (>90%) of excitatory glutamatergic synapses in the mammalian brain.
- Premisecitation
In the mouse visual cortex, depriving visual input from birth prevented the decrease in spine motility and maturation of spine morphology.
“In the mouse visual cortex, depriving visual input from birth prevented the decrease in spine motility and maturation of spine morphology (Majewska and Sur, 2003; Tropea et al., 2010).”
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- cites Tropea et al., 2010 source
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Only a small fraction of transient spines represent short-lived synaptic contacts.
“While stable spines mostly represent synaptic contacts, only a small fraction of transient spines represent short-lived synaptic contacts, and the rest of them represent failed synaptogenesis (Trachtenberg et al., 2002; Knott et al., 2006; Cane et al., 2014).”
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- cites Knott et al., 2006 source
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- cites Cane et al., 2014 source
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- The authors state this as an assumption: In principle, spine dynamics represent synapse dynamics.
- Premisecitation
Pharmacological blockade of NMDA receptors mimicked the effect of whisker trimming.
“Pharmacological blockade of NMDA receptors mimicked the effect of whisker trimming, indicating the involvement of the NMDA receptor pathway in such activity-dependent spine elimination (Zuo et al., 2005b).”
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- Premisecitation
The rest of transient spines represent failed synaptogenesis.
“While stable spines mostly represent synaptic contacts, only a small fraction of transient spines represent short-lived synaptic contacts, and the rest of them represent failed synaptogenesis (Trachtenberg et al., 2002; Knott et al., 2006; Cane et al., 2014).”
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- cites Cane et al., 2014 source
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- The authors state this as an assumption: In principle, spine dynamics represent synapse dynamics.
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Spine density varies significantly across diverse populations of neurons.
“Spine density varies significantly across diverse populations of neurons, likely reflecting the diversity of neuronal morphology and function (Nimchinsky et al., 2002; Ballesteros-Yanez et al., 2006).”
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Later on, sensory experience drives spine pruning (net loss of spines).
“Later on, sensory experience drives spine pruning (defined as net loss of spines).”
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The rates of spine formation and elimination change over time, resulting in non-monotonic alteration in spine density.
“The rates of spine formation and elimination change over time, resulting in non-monotonic alteration in spine density (Figure 1).”
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In the cerebral cortex, dendritic branches are mostly stable over time.
“In the cerebral cortex, while dendritic branches are mostly stable over time (Trachtenberg et al., 2002; Mizrahi and Katz, 2003; Chow et al., 2009; Mostany and Portera-Cailliau, 2011; Schubert et al., 2013), spines are constantly formed and eliminated.”
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Genetic deletion of the PirB receptor mimicked the effect of monocular deprivation on spine motility.
“Genetic deletion of the PirB receptor mimicked the effect of monocular deprivation on spine motility (Djurisic et al., 2013).”
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After initial net spine gain, spine elimination starts to outpace formation, leading to overall reduction of spine density.
“After this initial phase of net spine gain, spine elimination starts to outpace formation, leading to an overall reduction of spine density (Holtmaat et al., 2005; Zuo et al., 2005b; Yang et al., 2009).”
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Spine density continuously increases over P7-24 in rodent barrel cortex.
“Nevertheless, spine density continuously increases over this period of time (Cruz-Martin et al., 2010).”
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In adult animals, spine formation and elimination reach equilibrium.
“Finally, in adult animals spine formation and elimination reach equilibrium; spine density remains roughly constant until the onset of aging (Zuo et al., 2005a; Mostany et al., 2013).”
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- cites Mostany et al., 2013 source
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Spine density provides an approximate estimate of the total number of excitatory synapses onto the postsynaptic neuron.
“While spine density provides an approximate estimate of the total number of excitatory synapses onto the postsynaptic neuron, the location of a spine influences the contribution of its synaptically transmitted electrical and chemical signals to the integrated response at the soma (Nevian et al., 2007; Spruston, 2008).”
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In mice previously subjected to visual deprivation, light-induced spine maturation could be partially mimicked by pharmacological activation of the GABAergic system.
“In mice that had been subjected to visual deprivation previously, light-induced spine maturation could be partially mimicked by pharmacological activation of the GABAergic system, suggesting an important role of inhibitory circuits in the maturation of excitatory synapses (Tropea et al., 2010).”
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- Interpretationreasoning
Stronger synapses are more stable.
“As spine head size correlates with synaptic strength, this phenomenon suggests that stronger synapses are more stable (Holtmaat et al., 2005).”
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The location of a spine influences the contribution of its synaptically transmitted electrical and chemical signals to the integrated response at the soma.
“While spine density provides an approximate estimate of the total number of excitatory synapses onto the postsynaptic neuron, the location of a spine influences the contribution of its synaptically transmitted electrical and chemical signals to the integrated response at the soma (Nevian et al., 2007; Spruston, 2008).”
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Spine density remains roughly constant until the onset of aging in adult animals.
“Finally, in adult animals spine formation and elimination reach equilibrium; spine density remains roughly constant until the onset of aging (Zuo et al., 2005a; Mostany et al., 2013).”
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- cites Mostany et al., 2013 source
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- Interpretationcitation
The NMDA receptor pathway is involved in activity-dependent spine elimination.
“Pharmacological blockade of NMDA receptors mimicked the effect of whisker trimming, indicating the involvement of the NMDA receptor pathway in such activity-dependent spine elimination (Zuo et al., 2005b).”
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- Premisecitation
New spines were preferentially added onto layer 5 pyramidal neurons with complex apical tufts, rather than those with simple tufts.
“New spines were preferentially added onto layer 5 pyramidal neurons with complex apical tufts, rather than those with simple tufts (Holtmaat et al., 2006).”
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Spines on apical dendrites of layer 2/3 pyramidal neurons in rodent barrel cortex exhibit gradually decreasing motility and turnover rate between postnatal day 7 and 24.
“For example, spines on the apical dendrites of layer 2/3 pyramidal neurons in rodent barrel cortex exhibit gradually decreasing motility (elongation and shortening of spines) and turnover rate (defined as the total amount of gains and losses of spines) between postnatal day 7 and 24 (P7-24; Lendvai et al., 2000; Cruz-Martin et al., 2010).”
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The variation in spine density across neuronal populations likely reflects the diversity of neuronal morphology and function.
“Spine density varies significantly across diverse populations of neurons, likely reflecting the diversity of neuronal morphology and function (Nimchinsky et al., 2002; Ballesteros-Yanez et al., 2006).”
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Inhibitory circuits play an important role in the maturation of excitatory synapses.
“In mice that had been subjected to visual deprivation previously, light-induced spine maturation could be partially mimicked by pharmacological activation of the GABAergic system, suggesting an important role of inhibitory circuits in the maturation of excitatory synapses (Tropea et al., 2010).”
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- Premisecitation
In the cerebral cortex, spines are constantly formed and eliminated.
“In the cerebral cortex, while dendritic branches are mostly stable over time (Trachtenberg et al., 2002; Mizrahi and Katz, 2003; Chow et al., 2009; Mostany and Portera-Cailliau, 2011; Schubert et al., 2013), spines are constantly formed and eliminated.”
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During the early postnatal period, sensory inputs play instructive roles in the stabilization and maturation of spines.
“During early postnatal period, sensory inputs play instructive roles in the stabilization and maturation of spines.”
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The majority of stable spines formed before adolescence remain incorporated in the adult neuronal circuit.
“and the majority of stable spines formed before adolescence remain incorporated in the adult neuronal circuit (Zuo et al., 2005a; Yang et al., 2009; Yu et al., 2013).”
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Different motor skills are likely encoded by different subpopulations of synapses in the motor cortex.
“Furthermore, different motor skills are likely encoded by different subpopulations of synapses in the motor cortex, as learning a novel motor task in pre-trained mice continues to induce robust turnover in the adult motor cortex (Xu et al., 2009).”
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There exists a critical period during which surviving peri-infarct cortical tissues are most amenable to therapeutic interventions.
“This phenomenon suggests the existence of a critical period during which the surviving peri-infarct cortical tissues are most amenable to therapeutic interventions (Brown et al., 2007, 2009).”
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- Premisecitation
Following rapid spinogenesis during motor learning, spine density reverts to baseline level through elevated spine elimination.
“Following this rapid spinogenesis, spine density reverts to the baseline level through elevated spine elimination (Xu et al., 2009; Yu and Zuo, 2011).”
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Brief monocular deprivation increases spine formation along apical dendritic tufts of layer 5 pyramidal neurons in the binocular zone of the mouse visual cortex.
“Thus similar to chessboard trimming, MD has been found to increase spine formation along apical dendritic tufts of layer 5 pyramidal neurons in the binocular zone of the mouse visual cortex.”
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In a mouse model of Fragile X syndrome, spines are more numerous.
“In a mouse model of Fragile X syndrome, spines are more numerous, and a higher percentage of them appear immature upon examination of adult fixed tissues (Comery et al., 1997; Irwin et al., 2000).”
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A fear conditioning paradigm that pairs auditory cues with foot shocks has demonstrated opposite effects in auditory and frontal cortex.
“For example, a fear conditioning paradigm that pairs auditory cues with foot shocks has demonstrated opposite effects in auditory and frontal cortex.”
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The glucocorticoid level impacts motor learning-induced spine dynamics.
“Recently, it has also been found that the glucocorticoid level impacts motor learning-induced spine dynamics.”
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Alterations in dendritic spine densities have been observed in various neurological and neuropsychiatric diseases.
“Alterations in dendritic spine densities have been observed in various neurological and neuropsychiatric diseases.”
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In stroke models, severe ischemia leads to rapid spine loss, which is reversible after reperfusion if rescue is performed within 20-60 min.
“In stroke models, it is shown that severe ischemia leads to rapid spine loss, which is reversible after reperfusion if the rescue is performed within a short period of time (20–60 min; Zhang et al., 2005).”
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Survival of new spines correlates with retention of the motor skill.
“In mice, the amount of spines gained during initial learning closely correlates with the motor performance of learning acquisition (Xu et al., 2009); and survival of new spines correlates with retention of the motor skill (Yang et al., 2009).”
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- Interpretationcitation
Spine dynamics in different cortical regions may vary during the same task.
“More interestingly, spine dynamics in different cortical regions may vary during the same task.”
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In song birds, higher baseline spine turnover rate before song learning correlates with a greater capacity for subsequent song imitation.
“In song birds, higher baseline spine turnover rate before song learning has been found to correlate with a greater capacity for subsequent song imitation (Roberts et al., 2010).”
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The pattern of neural activity, rather than the magnitude, determines the stability of dendritic spines.
“Recently, an elegant study combining optogenetic stimulation and in vivo imaging showed that it is the pattern of neural activity, rather than the magnitude, that determines the stability of dendritic spines (Wyatt et al., 2012).”
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Glucocorticoid troughs following training were necessary for stabilization of spines formed during training and long-term memory retention.
“Training mice at glucocorticoid peaks resulted in higher rate of spine formation, whereas glucocorticoid troughs following training was necessary for stabilization of spines formed during training and long-term memory retention (Liston et al., 2013).”
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- Interpretationcitation
New spines formed during the initial monocular deprivation had functional synapses that were reactivated during the second monocular deprivation.
“Interestingly, a second MD failed to increase spine formation further, but selectively enlarged the spines formed during the initial MD, suggesting that new spines formed during the initial MD had functional synapses that were reactivated during the second MD (Hofer et al., 2009).”
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In an animal model of Huntington's disease, spine formation rate increases.
“In an animal model of Huntington’s disease spine formation rate increases, but newly formed spines do not persist to be incorporated into the local circuitry, which results in a net decrease in spine density (Murmu et al., 2013).”
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Tetrodotoxin blockade abolishes the effects of partial sciatic nerve ligation on spine remodeling.
“Such effects could be abolished by tetrodotoxin blockade, indicating that post-lesion spine remodeling is activity-dependent (Kim and Nabekura, 2011).”
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In mice, the amount of spines gained during initial learning closely correlates with the motor performance of learning acquisition.
“In mice, the amount of spines gained during initial learning closely correlates with the motor performance of learning acquisition (Xu et al., 2009); and survival of new spines correlates with retention of the motor skill (Yang et al., 2009).”
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The amount of new persistent spines after initial cocaine exposure correlated with the preference for the cocaine-paired context.
“Using a cocaine-conditioned place preference paradigm, a recent imaging study showed that initial cocaine exposure promoted spine formation in the frontal cortex, and that the amount of new persistent spines correlated with the preference for the cocaine-paired context (Munoz-Cuevas et al., 2013).”
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In the mouse motor cortex, spine formation begins immediately as the animal learns a new task.
“In the mouse motor cortex, spine formation begins immediately as the animal learns a new task.”
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In αCaMKII-T286A defective mice, chessboard trimming failed to increase stabilization of new persistent spines at the border between spared and deprived barrels.
“In αCaMKII-T286A defective mice, chessboard trimming failed to increase stabilization of new persistent spines at the border between spared and deprived barrels (Wilbrecht et al., 2010).”
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In an animal model of Huntington's disease, newly formed spines do not persist to be incorporated into the local circuitry.
“In an animal model of Huntington’s disease spine formation rate increases, but newly formed spines do not persist to be incorporated into the local circuitry, which results in a net decrease in spine density (Murmu et al., 2013).”
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In an animal model of Huntington's disease, there is a net decrease in spine density.
“In an animal model of Huntington’s disease spine formation rate increases, but newly formed spines do not persist to be incorporated into the local circuitry, which results in a net decrease in spine density (Murmu et al., 2013).”
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Initial cocaine exposure promoted spine formation in the frontal cortex.
“Using a cocaine-conditioned place preference paradigm, a recent imaging study showed that initial cocaine exposure promoted spine formation in the frontal cortex, and that the amount of new persistent spines correlated with the preference for the cocaine-paired context (Munoz-Cuevas et al., 2013).”
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Altered spine dynamics has been reported in animal models of degenerative diseases.
“Altered spine dynamics has also been reported in animal models of degenerative diseases.”
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In vivo imaging studies have shown that in the cerebral cortex, spine dynamics directly correlates with learning.
“Indeed, in vivo imaging studies have shown that in the cerebral cortex, spine dynamics directly correlates with learning.”
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In the auditory cortex, increased spine formation was correlated with paired fear conditioning.
“In the auditory cortex, it was found that increased spine formation was correlated with paired fear conditioning, while unpaired conditioning was associated with increased elimination of spines (Moczulska et al., 2013).”
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- Interpretationcitation
Post-lesion spine remodeling is activity-dependent.
“Such effects could be abolished by tetrodotoxin blockade, indicating that post-lesion spine remodeling is activity-dependent (Kim and Nabekura, 2011).”
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- Premisecitation
In the auditory cortex, unpaired conditioning was associated with increased elimination of spines.
“In the auditory cortex, it was found that increased spine formation was correlated with paired fear conditioning, while unpaired conditioning was associated with increased elimination of spines (Moczulska et al., 2013).”
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Addiction elicits similar temporal changes in spine dynamics as motor learning does.
“Addiction, which has been considered as pathological learning (Hyman, 2005), elicits similar temporal changes in spine dynamics as motor learning does.”
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Spine loss is accelerated in the vicinity of β-amyloid plaques in the cerebral cortex.
“For example, spine loss is accelerated in the vicinity of β-amyloid plaques in the cerebral cortex (Tsai et al., 2004; Spires et al., 2005).”
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There is growing consensus that spine abnormality is associated with behavioral deficiency and decline in cognitive functions.
“There is growing consensus that spine abnormality is associated with behavioral deficiency and decline in cognitive functions (for detail see Fiala et al., 2002; Penzes et al., 2011).”
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In a mouse model for chronic pain, partial sciatic nerve ligation increases spine formation and elimination.
“In a mouse model for chronic pain, partial sciatic nerve ligation increases spine formation and elimination.”
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- Interpretationreasoning
These studies reveal diversity of temporal rules underlying learning-induced spine dynamics.
“Taken together, these studies reveal the diversity of temporal rules underlying learning-induced spine dynamics.”
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Whether spines are formed or removed during learning depends on the behavioral paradigm as well as the specific neuronal circuit and cell types participating in the learning process.
“Whether spines are formed or removed during learning depends on the behavioral paradigm as well as the specific neuronal circuit and cell types participating in the learning process.”
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Training mice at glucocorticoid peaks resulted in a higher rate of spine formation.
“Training mice at glucocorticoid peaks resulted in higher rate of spine formation, whereas glucocorticoid troughs following training was necessary for stabilization of spines formed during training and long-term memory retention (Liston et al., 2013).”
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Monocular deprivation significantly increases coordinated dynamics of spines and inhibitory synapses nearby in layer 2/3 pyramidal neurons.
“Monocular deprivation significantly increases coordinated dynamics of spines and the inhibitory synapses nearby in layer 2/3 pyramidal neurons (Chen et al., 2012).”
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- Interpretationcitation
These findings support the clustered plasticity model, which postulates that clustered synapses are more likely to participate in encoding the same information than synapses dispersed throughout the dendritic arbor.
“These findings support the clustered plasticity model, which postulates that clustered synapses are more likely to participate in encoding the same information than synapses dispersed throughout the dendritic arbor (Govindarajan et al., 2006).”
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In the fear conditioning paradigm, a spine eliminated during fear conditioning is usually replaced by a spine in its vicinity (within 2 μm) during fear extinction.
“Similar spatial selectivity of spine dynamics has been observed in the fear conditioning paradigm: a spine eliminated during fear conditioning is usually replaced by a spine in its vicinity (within 2 μm) during fear extinction (Lai et al., 2012).”
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In mice overexpressing MECP2, both spine gains and losses are elevated.
“In mice overexpressing MECP2, a Rett Syndrome related gene, it has been found that both spine gains and losses are elevated.”
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In the mouse auditory cortex, spines tuned for different peak frequencies are interspersed along the dendrites of pyramidal neurons.
“Combining in vivo whole-cell patch recording and single spine calcium imaging, a recent work has shown that spines tuned for different peak frequencies are interspersed along dendrites of pyramidal neurons in the mouse auditory cortex (Chen et al., 2011).”
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The emergence and disappearance of spines are neither uniform nor random along dendrites, but rather occur at spatially selective 'hot spots'.
“Structural imaging of spines has suggested that the emergence and disappearance of spines are neither uniform nor random along dendrites, but rather occur at spatially selective “hot spots.””
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Microglial processes and spine contact are involved in spine elimination.
“The motility of microglial processes and spine contact are actively regulated by sensory experience and are involved in spine elimination (Tremblay et al., 2010).”
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Addition of the second new spine in a cluster is often associated with enlargement of the first new spine.
“Furthermore, addition of the second new spine in the cluster is often associated with the enlargement of the first new spine.”
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Spine dynamics are influenced by dynamics of inhibitory synapses.
“Interestingly, spine dynamics are also influenced by dynamics of inhibitory synapses.”
Quote in the paperNot a substantial claim - Premisecitation
Selective removal of BDNF in microglia recapitulates the effects of microglial depletion on motor learning induced spine formation.
“In addition, the depletion of microglia resulted in significant reduction of motor learning induced spine formation, and selective removal of brain-derived neurotrophic factor (BDNF) in microglia recapitulated the effects of microglial depletion (Parkhurst et al., 2013).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Parkhurst et al., 2013 source
Not checked in the free run.
- Premisecitation
Microglia are in close contact with dendritic spines.
“Another type of glial cells, microglia, have also been found to be in close contact with dendritic spines.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Tremblay et al., 2010 source
Not checked in the free run.
- Interpretationreasoning
Repeated re-activation of the first new spine is required for the clustered emergence of the second new spine.
“Taken together, these observations suggest that repeated re-activation of the first new spine is required for the clustered emergence of the second new spine.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Fu et al., 2012
Not checked in the free run.
- Premisecitation
Depletion of microglia results in significant reduction of motor learning induced spine formation.
“In addition, the depletion of microglia resulted in significant reduction of motor learning induced spine formation, and selective removal of brain-derived neurotrophic factor (BDNF) in microglia recapitulated the effects of microglial depletion (Parkhurst et al., 2013).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Parkhurst et al., 2013 source
Not checked in the free run.
- Premisecitation
In Fragile X syndrome model animals, spine turnover increased in various cortical areas.
“In vivo studies further showed that in such animals spine turnover increased in various cortical areas (Cruz-Martin et al., 2010; Pan et al., 2010; Padmashri et al., 2013), and neither whisker trimming nor motor learning could further alter spine dynamics (Pan et al., 2010; Padmashri et al., 2013).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Premisecitation
Blockade of astrocytic glutamate uptake accelerates experience-dependent spine elimination during adolescent development.
“For example, blockade of astrocytic glutamate uptake has been shown to accelerate experience-dependent spine elimination during adolescent development (Yu et al., 2013).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Yu et al., 2013
Not checked in the free run.
- Premisecitation
Spines formed during tandem execution of different motor tasks or during motor enrichment do not cluster.
“In contrast, spines formed during tandem execution of different motor tasks or during motor enrichment do not cluster (Fu et al., 2012).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Fu et al., 2012
Not checked in the free run.
- Premisecitation
In a mouse model of Fragile X syndrome, a higher percentage of spines appear immature upon examination of adult fixed tissues.
“In a mouse model of Fragile X syndrome, spines are more numerous, and a higher percentage of them appear immature upon examination of adult fixed tissues (Comery et al., 1997; Irwin et al., 2000).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Premisecitation
In the mouse motor cortex, new spines that form during repeated training with the same motor task tend to cluster.
“In the mouse motor cortex, new spines that form during repeated training with the same motor task tend to cluster.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Fu et al., 2012
Not checked in the free run.
- May combine two claims. 17
- Classified as probably not a substantial claim. 4
May combine two claims.
PremisecitationSpine morphology and dynamics vary among neuronal types, across developmental stages, and in response to experiences such as sensory stimulation and deprivation, environmental enrichment, and various paradigms of learning.
“Furthermore, their morphology and dynamics vary among neuronal types, across developmental stages, and in response to experiences such as sensory stimulation and deprivation, environmental enrichment, and various paradigms of learning (Holtmaat and Svoboda, 2009; Fu and Zuo, 2011).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
Premisenone givenBoth acute and chronic sensory manipulations profoundly impact spine dynamics, but the exact effect depends on the manipulation paradigm and duration, as well as the developmental stage of the animal.
“Both acute and chronic sensory manipulations have been shown to profoundly impact spine dynamics, but the exact effect depends on the manipulation paradigm and duration, as well as the developmental stage of the animal.”
Quote in the paperSubstantial claimMay combine two claims.
PremisereasoningSpines form, enlarge, shrink, and retract throughout the animal's lifespan.
“From such time-lapse imaging studies a dynamic picture of spines has emerged: spines form, enlarge, shrink, and retract throughout the animal’s lifespan.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationUnilateral trimming of all whiskers in 1-month-old mice for 4 or 14 days dramatically reduced spine elimination in the barrel cortex, but left spine formation largely unperturbed.
“Unilateral trimming of all whiskers in 1-month-old mice for 4 or 14 days dramatically reduced spine elimination in the barrel cortex, but left spine formation largely unperturbed (Zuo et al., 2005b; Yu et al., 2013).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Zuo et al., 2005b
Not checked in the free run.
- cites Yu et al., 2013
Not checked in the free run.
May combine two claims.
PremisecitationNot all spines are equally susceptible to elimination: those with large heads are more stable than thin ones.
“Importantly, not all spines are equally susceptible to elimination: those with large heads are more stable than thin ones.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Holtmaat et al., 2005 source
Not checked in the free run.
May combine two claims.
PremisecitationChessboard trimming promotes spine turnover and stabilizes newly formed spines selectively in a subclass of cortical neurons.
“Such paradigm has been shown to promote spine turnover and to stabilize newly formed spines selectively in a subclass of cortical neurons (Trachtenberg et al., 2002; Holtmaat et al., 2006).”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationBetween P28 and P42, 17% of spines are eliminated along apical dendrites of layer 5 pyramidal neurons in mouse barrel cortex, while only 5% of new spines are formed during the same period.
“Between P28 and P42, 17% of spines are eliminated along the apical dendrites of layer 5 pyramidal neurons in the mouse barrel cortex, while only 5% of new spines are formed during the same period of time (Zuo et al., 2005a, b).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Zuo et al., 2005a, b
Not checked in the free run.
Classified as probably not a substantial claim.
InterpretationreasoningSpines may serve as the structural substrate for learning and memory.
“The highly dynamic nature of dendritic spines elicits the prevalent idea that spines may serve as the structural substrate for learning and memory.”
Quote in the paperSubstantial claim- The authors state this as an assumption: In principle, spine dynamics represent synapse dynamics.
May combine two claims.
PremisecitationIn the chronic pain model, elevation of spine formation rate precedes that of elimination, leading to an initial increase in spine density followed by its reduction.
“Similar to the stroke model, elevation of spine formation rate precedes that of elimination, leading to an initial increase in spine density followed by its reduction.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Kim and Nabekura, 2011
Not checked in the free run.
May combine two claims.
PremisecitationA second monocular deprivation failed to increase spine formation further, but selectively enlarged the spines formed during the initial monocular deprivation.
“Interestingly, a second MD failed to increase spine formation further, but selectively enlarged the spines formed during the initial MD, suggesting that new spines formed during the initial MD had functional synapses that were reactivated during the second MD (Hofer et al., 2009).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Hofer et al., 2009 source
Not checked in the free run.
May combine two claims.
PremisecitationInjury-induced plasticity after stroke reaches its peak at 1 week post-stroke; from then on the rate of spine formation and elimination steadily decline.
“This injury-induced plasticity reaches its peak at 1 week post-stroke; from then on the rate of spine formation and elimination steadily decline.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Brown et al., 2007, 2009
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
PremisecitationWhile neurodegenerative diseases are usually associated with net spine loss, neurodevelopmental disorders exhibit diverse spine phenotypes.
“While neurodegenerative diseases are usually associated with net spine loss, neurodevelopmental disorders exhibit diverse spine phenotypes.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationFollowing stroke, spine formation and subsequent elimination increase in the peri-infarct region, but not in cortical territories distant from the infarct or in the contralateral hemisphere.
“Following stroke, spine formation and subsequent elimination increase in the peri-infarct region, but not in cortical territories distant from the infarct or in the contralateral hemisphere (Brown et al., 2009; Johnston et al., 2013).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Brown et al., 2009 source
Not checked in the free run.
- cites Johnston et al., 2013
Not checked in the free run.
May combine two claims.
PremisecitationThe effect of monocular deprivation on spine formation was not observed in layer 2/3 neurons or in the monocular zone.
“However this effect was not observed in layer 2/3 neurons, or in the monocular zone (Hofer et al., 2009), again indicating a cell type specific synapse remodeling.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Hofer et al., 2009 source
Not checked in the free run.
May combine two claims.
InterpretationreasoningEach disorder presents with its own hallmark abnormalities in spine dynamics, which corroborates the idea that spines are structural underpinnings for proper cognitive functioning.
“Each disorder presents with its own hallmark abnormalities in spine dynamics, which further corroborates the idea that spines are structural underpinnings for proper cognitive functioning.”
Quote in the paperNot a substantial claim- The authors state this as an assumption: In principle, spine dynamics represent synapse dynamics.
May combine two claims.
PremisecitationThe motility of microglial processes and spine contact are actively regulated by sensory experience.
“The motility of microglial processes and spine contact are actively regulated by sensory experience and are involved in spine elimination (Tremblay et al., 2010).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Tremblay et al., 2010 source
Not checked in the free run.
Classified as probably not a substantial claim.
Premisenone givenAll the works discussed in this review have focused on the postsynaptic side of the synapse.
“It is worth noting that all works discussed above have focused on the postsynaptic side, which is only half of the story.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationIn Fragile X syndrome model animals, neither whisker trimming nor motor learning could further alter spine dynamics.
“In vivo studies further showed that in such animals spine turnover increased in various cortical areas (Cruz-Martin et al., 2010; Pan et al., 2010; Padmashri et al., 2013), and neither whisker trimming nor motor learning could further alter spine dynamics (Pan et al., 2010; Padmashri et al., 2013).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Pan et al., 2010 source
Not checked in the free run.
- cites Padmashri et al., 2013
Not checked in the free run.
Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing), S4 (based).
Premisenone givenThe most intriguing role of glial cells is their participation at synaptic functioning and dynamics.
“The most intriguing role of glial cells is their participation at synaptic functioning and dynamics.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationIn mice overexpressing MECP2, new spines are more vulnerable to elimination than in wild type mice, resulting in a net loss of spines.
“However, new spines are more vulnerable to elimination than in wild type mice, resulting in a net loss of spines (Jiang et al., 2013).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Jiang et al., 2013 source
Not checked in the free run.
Classified as probably not a substantial claim.
PremisereasoningIdentification of the presynaptic partner of an imaged dendritic spine remains a technical challenge, because the presynaptic axon may originate from many sources and is usually intermingled with other axonal processes.
“However, identification of the presynaptic partner of an imaged dendritic spine remains a technical challenge, as the presynaptic axon may originate from a plethora of sources, and is usually intermingled with many other axonal processes.”
Quote in the paperSubstantial claim
The paper relies on this without support.
Dendritic spines are the postsynaptic sites of the great majority (>90%) of excitatory glutamatergic synapses in the mammalian brain.
“They are the postsynaptic sites of the great majority (>90%) of excitatory glutamatergic synapses in the mammalian brain, and contain essential molecular components for postsynaptic signaling and plasticity.”
The authors state this as an assumption.
In principle, spine dynamics represent synapse dynamics.
“In principle, spine dynamics represent synapse dynamics.”
The paper relies on this without support.
The cerebral cortex has the ability to reorganize its circuitry in response to experiences.
“The cerebral cortex has the amazing ability to reorganize its circuitry in response to experiences.”
The authors state this as an assumption.
Chessboard trimming amplifies differences in activity levels and patterns of neighboring barrels, thereby introducing a novel sensory experience.
“While complete whisker trimming removes sensory input globally, trimming every other whisker (“chessboard trimming”) presumably amplifies any difference in activity levels and patterns of neighboring barrels, thereby introducing a novel sensory experience.”
The paper relies on this without support.
Brief monocular deprivation increases the disparity between inputs from the two eyes.
“Similar to chessboard trimming, brief monocular deprivation (MD) increases the disparity between the inputs from two eyes.”
The paper relies on this without support.
The hippocampus is buried beneath the cortex and beyond the reach of standard two-photon microscopy.
“On one hand, hippocampus, the structure crucial for formation of declarative memory, is buried beneath cortex and beyond the reach of standard two-photon microscopy.”
The paper relies on this without support.
The presynaptic side (the identity and geometry of presynaptic axons and the availability of axonal boutons) is a major determinant of spine distribution and dynamics.
“The other major determinant of spine distribution and dynamics lies at the presynaptic side: the identity and geometry of presynaptic axons and the availability of axonal boutons.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Across the mouse neocortex, rates of spine gain and spine loss decrease from peripubertal age (P30) to P60.
“Across the mouse neocortex, rates of spine gain and spine loss (independent of spine density) decrease from peripubertal age (P30) to P60 (Chen et al., 2014; Johnson et al., 2016a; Zuo et al., 2005).”
From Does puberty mark a transition in sensitive periods for plasticity in the associative neocortex? (2017) · cites this paper as Chen et al., 2014
The citation lists haven’t been fetched yet
References 71 · Cited by 88
- Authors
- Chen, C.-C., Lu, J., & Zuo, Y.
- Affiliations
- University of California, Santa Cruz (US)
- Stanford University (US)
- Corresponding author
- Zuo, Yi
- Venue
- Frontiers in Neuroanatomy
- Publisher
- Frontiers Media SA
- Published
- 2014-05-09
- Volume
- 8
- Pages
- 28-28
- ISSN
- 1662-5129 (ISSN-L)
- PMC
- PMC4023020
- OpenAlex
- W2141331098
- Semantic Scholar
- 09a8a143996cb58519d4a68f10700f1632bc144b
- Cited by
- 88
- References
- 71
- Citations by year
- 2024: 4 · 2023: 5 · 2022: 11 · 2021: 11 · 2020: 4 · 2019: 8 · 2018: 11 · 2017: 13 · 2016: 11 · 2015: 8 · 2014: 2
- FWCI
- 1.29
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- cc-by
- Locations
- Frontiers in Neuroanatomy (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- eScholarship (California Digital Library) (submittedVersion)
- Funders
- National Institute of Mental Health (R01MH094449)
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetics and Neurodevelopmental Disorders
- Keywords
- dendritic spines, spine dynamics, postsynaptic sites, excitatory synapses, synaptogenesis, synaptic pruning, memory encoding, in vivo imaging, spatiotemporal patterns, brain development, neurodevelopmental disorders
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