Table 2.
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 |
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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) |