Neuronal activation in female budgerigars is localized and related to male song complexity

Females of several songbird species have been shown to respond preferentially to a more complex song. The male budgerigar (Melopsittacus undulatus) sings complex songs consisting of discrete components, known as syllables. We exposed female budgerigars to either standard male song, complex song, or simple song, the iteration of only one syllable (either frequency-modulated or unmodulated). Using immunocytochemistry, we analysed the expression of the protein product of the immediate early gene ZENK in a number of forebrain regions. The level of Zenk protein expression caused by song stimuli varied among each of the brain regions. Expression was highest in the caudomedial neostriatum (NCM), lower in the caudomedial hyperstriatum ventrale (CMHV), and lowest in the hippocampus. There was a significant effect of song complexity on the number of Zenk-immunoreactive cells in the NCM, but not in the hippocampus. Zenk protein expression correlated significantly and positively with the number of different syllables to which the females were exposed in the NCM and to a lesser extent in the CMHV, but not in the hippocampus. For the NCM this correlation was also significant within the group exposed to natural song. These results suggest that the NCM is involved in the perception of song complexity in female budgerigars.