Impact of experience-dependent and -independent factors on gene expression in songbird brain.

Songbirds provide rich natural models for studying the relationships between brain anatomy, behavior, environmental signals, and gene expression. Under the Songbird Neurogenomics Initiative, investigators from 11 laboratories collected brain samples from six species of songbird under a range of experimental conditions, and 488 of these samples were analyzed systematically for gene expression by microarray. ANOVA was used to test 32 planned contrasts in the data, revealing the relative impact of different factors.

LMAN lesions prevent song degradation after deafening without reducing HVC neuron addition.

In some songbirds perturbing auditory feedback can promote changes in song structure well beyond the end of song learning. One factor that may drive vocal change in such deafened birds is the ongoing addition of new vocal-motor neurons into the song system. Without auditory feedback to guide their incorporation, the addition of these new neurons could disrupt the established song pattern.

Song tutoring triggers CaMKII phosphorylation within a specialized portion of the avian basal ganglia.

In several songbird species, a specialized anterior forebrain pathway (AFP) that includes part of the avian basal ganglia has been implicated specifically in song learning. To further elucidate cellular mechanisms and circuitry involved in vocal learning, we used quantitative immunoblot analysis to determine if early song tutoring promotes within the AFP phosphorylation of calcium/calmodulin-dependent kinase II (CaMKII), a multifunctional kinase whose phosphorylation at threonine 286 is critical for many forms of neural plasticity and behavioral learning. We report that in young male zebra finches likely to have begun the process of song acquisition, brief tutoring by a familiar conspecific adult promotes a dramatic increase in levels of phosphorylated CaMKII (pCaMKII) in Area X, the striatal/pallidal component of the AFP.

Synaptic and molecular mechanisms regulating plasticity during early learning.

Many behaviors are learned most easily during a discrete developmental period, and it is generally agreed that these "sensitive periods" for learning reflect the developmental regulation of molecular or synaptic properties that underlie experience-dependent changes in neural organization and function. Avian song learning provides one example of such temporally restricted learning, and several features of this behavior and its underlying neural circuitry make it a powerful model for studying how early experience sculpts neural and behavioral organization. Here we describe evidence that within the basal ganglia-thalamocortical loop implicated in vocal learning, song acquisition engages N-methyl-d-aspartate receptors (NMDARs), as well as signal transduction cascades strongly implicated in other instances of learning.

Developmental patterns of NMDAR expression within the song system do not recur during adult vocal plasticity in zebra finches.

All songbirds learn to sing during postnatal development but then display species differences in the capacity to learn song in adulthood. While the mechanisms that regulate avian vocal plasticity are not well characterized, one contributing factor may be the composition of N-methyl-D-aspartate receptors (NMDAR). Previous studies of an anterior forebrain pathway implicated in vocal plasticity revealed significant regulation of NMDAR subunit expression during the developmental sensitive period for song learning.