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  • To determine whether spine dynamics were affected in ephrin-A KOs, we repeatedly imaged apical dendritic branches and followed spine dynamics in the motor cortex by transcranial two-photon microscopy. We found that, while the amount of new spines added over 2?days was comparable between ephrin-A2 Alisertib price KOs and their wild-type littermates, significantly more spines were eliminated during the same period of time in KOs ( Figures 1A, 1B, and 1E; p?< 0.001). Unlike ephrin-A2 KOs, ephrin-A3 KOs exhibited similar spine turnover to wild-type controls ( Figures 1C and 1E; p > 0.1). In addition, spine elimination in ephrin-A2/A3 double KOs was comparable to that of ephrin-A2 single KOs ( Figures 1D and 1E; p > 0.7). Moreover, we found that the increase in spine elimination occurred in various cortical regions of ephrin-A2 KOs ( Figure?S2) and persisted over prolonged imaging intervals ( Figure?1F). As a consequence, despite the normal spine density at 1?month of age (p > 0.2), spine Cobimetinib cost density of adult ephrin-A2 KOs was lower than that of wild-type mice ( Figure?1G; p?< 0.05). Thus, developmentally regulated spine pruning is accelerated in ephrin-A2 KOs. Spines formed during early development and surviving extensive pruning have greatly contributed to stably connected neural networks in adulthood (Yang et?al., 2009?and?Zuo et?al., 2005a). To investigate how accelerated spine elimination during adolescent development influences adult synaptic connections in ephrin-A2 KOs, we calculated the lifetime of dendritic spines in both wild-type and ephrin-A2 KO mice, based on spine elimination measured over various imaging intervals (i.e., 2, 4, 8, 30, and 90?days), starting at 1?month of age ( Figure?1F). We found that the spine survival curve of wild-type mice was well fitted by a two-phase exponential decay equation (R2?= 0.92, see Experimental Procedures), Bumetanide with a small portion of spines rapidly lost (��fast-decay spines,�� 15%, half-life 3.0?days) and the rest stable over months (��slow-decay spines,�� half-life 1,112?days). Fitting the spine survival curve of ephrin-A2 KOs with the same formula (R2?= 0.90), we found that while the half-life of the fast-decay spine population in KOs was comparable to that of wild-type mice (2.3?days, p > 0.4), the half-life of the slow-decay spine population was significantly shorter in KOs (432?days, p?< 0.01). As previous studies have revealed that newly formed spines are much more vulnerable to elimination than pre-existing spines ( Xu et?al., 2009?and?Yang et?al., 2009), the fast- and slow-decay spines could represent new and pre-existing spines, respectively. To determine whether the survival of new and pre-existing spines was, indeed, affected differently in ephrin-A2 KOs, we imaged the same mice three times (i.e.

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