“…Neurons in HVC, RA, Area X, and LMAN regulate various aspects of song control and have been implicated in the social modulation of song (Sakata and Vehrencamp, 2012;Brainard and Doupe, 2013;Bertram et al, 2014;Woolley and Kao, 2015). For example, neurophysiological activity in Area X, LMAN, and RA is significantly different when male songbirds perform UD and FD song (Hessler and Doupe, 1999;Kao et al, 2005;Kao and Brainard, 2006;Kao et al, 2008;Sober et al, 2008;Woolley et al, 2014), and manipulations of Area X, LMAN, and RA activity significantly affect the control and social modulation of syllable structure (Vu et al, 1994;Ashmore et al, 2005Ashmore et al, , 2008€ Olveczky et al, 2005;Kao and Brainard, 2006;Hampton et al, 2009;Leblois et al, 2010;Stepanek and Doupe, 2010;Figure 6 Relationship between song features and EGR-1 expression in song control nuclei, independent of social context.…”