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. 2020 Aug 28;12:36. doi: 10.3389/fnsyn.2020.00036

Table 2.

Experience-dependent spine plasticity.

Modulation Brain region Age animals Impact on spine formation/elimination Main results Methods Reference
Whisker trimming
Chessboard whisker trimming Barrel cortex, PN, Layer 5 6-10 weeks Control: 40% stable spines over 4 days Trimmed: 30% stable spines over 4 days Sensory deprivation increases spine turnover and reduces stability. Two-photon laser scanning, Craniotomy, GFP-labeled dendrites Trachtenberg et al. (2002)
Unilateral whisker trimming Barrel cortex, PN, Layer 5 P30 Control: 17% of spines eliminated and 6% formed over 2 weeks Trimmed: 10% of spines eliminated, 5% formed after 2 weeks Long-term sensory deprivation in young mice reduces the rate of spine elimination but has no significant effect on spine formation. Spines in adulthood are less affected. Two-photon laser scanning, Thinned skull cranial window, YFP-labeled dendrites Zuo et al. (2005a)
>4 months Control: 5% of spines eliminated and 4% formed over 2 weeks Trimmed: No changes in spine turnover after 2 weeks
Chessboard whisker trimming Barrel cortex, PN, Layer 5 2-5 months Control: ~63% of spines stable over 28 days Trimmed: ~60% of spines stable over 28 days, turnover increased Sensory deprivation induces loss of old persistent spines and forms new persistent spines. Two-photon laser scanning, Craniotomy, GFP-labeled dendrites, Electron microscopy Holtmaat et al. (2006)
Motor learning
Motor task, Neonatal bilateral whisker trimming Barrel cortex, Motor cortex, PN, Layer 2/3, Layer 5 P30 Control, MC, L2/3: ~18% spine elimination, ~18% spine formation over 4 days Motor task, MC, L2/3: ~16% spine elimination, ~17% spine formation Motor task-induced increase in spine dynamics happens only in L5, but not in L2/3 of MC. Neonatal whisker trimming reduces spine formation in L2/3, but not in L5 of the somatosensory cortex. Two-photon laser scanning, Thinned skull cranial window, GFP-labeled dendrites Tjia et al. (2017)
Control, MC, L5: ~9% spine elimination, ~6% spine formation over 4 days
Motor task, MC, L5: ~14% spine elimination ~14% spine formation
Control, BC, L2/3: ~15% spine elimination, ~15% spine formation After neo. trimming, BC, L2/3: ~17% spine elimination, ~7% spine formation
Control, BC, L5: ~12% spine elimination, ~7% spine formation After neo. trimming, BC, L5: ~12% spine elimination, ~7% spine formation
Forelimb reaching Motor cortex, PN, L5 P30 Control: ~7% spine elimination, ~5% spine formation over 2 days Motor learning selectively stabilizes learning-induced new spines into adulthood. Two-photon laser scanning, Thinned skull cranial window and craniotomy, YFP-labeled dendrites Xu et al. (2009)
Reaching: Spine elimination increased after 2 days (~15%), spine formation increased to 11% within 1 h after training
Rotarod motor task Motor cortex, PN, Layer 5 P30 Control MC: ~9% elimination, ~7% spine formation over 2 days Rotarod MC: ~9% elimination, ~15% spine formation over 2 days Learning induces formation of new spines. Two-photon laser scanning, Thinned skull cranial window, YFP-labeled dendrites Yang et al. (2009)
>4 months Control MC: ~3% elimination, ~3% spine formation over 2 days Rotarod MC: ~4% elimination, ~8% spine formation over 2 days
Visual deprivation
Monocular deprivation Visual cortex, PN, Layer 2/3, Layer 5 P45-100 Control L2/3: 8% spine elimination, 7% spines formation over 8 days Control L5: 7% spine elimination, 6% spines formation over 4 days Visual deprivation increases spine formation. Two-photon laser scanning, Craniotomy, GFP- labeled dendrites Hofer et al. (2009)
MD L2/3: no changes in spine turnover over 4+ days MD L5: spine elimination unchanged, ~11% of spines formed over 4 days
Monocular- and Binocular deprivation Visual cortex, PN, Layer 5 P28 Control: ~11% spine elimination, ~8% spine formation over 3 days MD: ~19% spine elimination, ~9% spine formation over 3 days BD: ~10% spine elimination, ~7% spine formation over 3 days MD over 3 days significantly increases spine elimination without affecting spine formation. BD does not change spine dynamics. Two-photon laser scanning, Thinned skull cranial window, YFP-labeled dendrites Zhou et al. (2017)
Fear conditioning
Fear conditioning, Fear extinction Frontal association cortex, PN, Layer 5 P30 Control: ~18% spine elimination, ~14% spine formation over 9 days Fear cond.: ~23% spine elimination, ~11% spine formation over 9 days Fear ext.: ~10% spine elimination, ~17% spine formation after 2 days Fear conditioning promotes spine elimination. Fear extinction induces spine formation. Two-photon laser scanning, Thinned skull cranial window, YFP-labeled dendrites Lai et al. (2012)
Fear conditioning Auditory cortex, PN, Layer 5 3-6 months Control: ~7% spine elimination, ~8% spine formation over 2 h Fear cond.: ~11% spine elimination, ~17% spine formation over 2 h Auditory fear conditioning causes an increase of spine turnover Two-photon laser scanning, Craniotomy, GFP-labeled dendrites Lai et al. (2018)
Fear conditioning Auditory cortex, PN, Layer 5 7–10 weeks Control: ~13% spine elimination, ~7% spine formation over 3 days Fear cond.: ~13% spine elimination, ~15% spine formation Fear conditioning increases formation of new Amygdala–Auditory cortex connections consistent with the consolidation of fear memory. Two-photon laser scanning, Craniotomy, YFP, tdTomato and GFP-labeled dendrites and axons Yang et al. (2016)
Fear conditioning, Fear extinction Auditory cortex, PN, Layer 5 P30 Control: ~9% spine elimination, ~9% spine formation over 3 days Fear cond.: ~10% spine elimination, ~16% spine formation over 3 days Fear ext.: ~17% spine elimination, ~5% spine formation over 2 days Persistent new spines are induced by auditory fear conditioning. Fear extinction selectively eliminates new spines. Two-photon laser scanning, Thinned skull cranial window, YFP-labeled dendrites Lai et al. (2018)
Stress
Corticosterone administration (stress) Barrel cortex, PN, Layer 5 P23-30 Control: ~4% elimination, ~5% spine formation over 1 day Acute cort.: ~12% elimination, ~7% spine formation over 1 day Chronic cort.: Elimination increases to 22%, Spine formation unchanged over 10 days Acute corticosterone increases spine turnover. Chronic stress increases spine elimination. Two-photon laser scanning, Thinned skull cranial window, YFP-labeled dendrites Liston and Gan (2011)
Motor task, corticosterone administration (stress) Motor cortex, PN, Layer 5 P30 Untrained: ~7% spine formation over 2 days Training with additional cort: ~17% spine formation over 2 days Chronic cort.: elimination of training associated and pre-training spines over 10 days Corticosterone increases formation of lasting task-associated spines. Chronic corticosterone causes loss of spines and reduces motor performance. Two-photon laser scanning, Thinned skull cranial window, YFP-labeled dendrites Liston et al. (2013)