NR2B downregulation in a forebrain region required for avian vocal learning is not sufficient to close the sensitive period for song learning

The neural changes that limit the sensitive period for avian song development are unknown, but neurons in a forebrain region critical for song learning, the lMAN, exhibit experience-driven changes in NMDAR subunit expression that could regulate sensitive period closure. Specifically, NR2B levels in lMAN decrease during song acquisition, potentially reducing synaptic plasticity by decreasing NMDAR EPSC duration and/or affecting NMDAR-coupled intracellular cascades. While rearing birds in isolation extends the sensitive period and also delays the developmental changes in NR2B expression and NMDAR physiology, recent work indicates that a transition to faster NMDAR currents does not preclude further song learning. However, NR2B mRNA expression in isolates remains elevated beyond the age at which NMDAR currents shorten, leaving open the possibility that NR2B levels regulate closure of the sensitive period through effects other than those mediated by NMDAR current duration. To determine whether the experience-driven decrease in NR2B expression in lMAN closes the sensitive period, we promoted this change in gene expression either by treating isolation-reared zebra finches briefly with testosterone (T-isolates) or by allowing males limited access to conspecific song (pre-exposed isolates). We then assessed if these birds could acquire song from tutors after the normal close of the sensitive period. Despite a normal decline in NR2B expression, T-isolate and pre-exposed isolate birds learned tutor songs heard from d65-90, while normally reared birds did not. These findings suggest that the normal decline in NR2B expression with lMAN is not sufficient for sensitive period closure.