Multiscale integrated structural model of yeast NuA4. (A) High-resolution structures of Tra1 (5OJS), core Piccolo NuA4 (5J9U), Esa1 Tudor domain (2RO0), Arp4-Act1 dimer (5I9E), and Yaf9 YEATS domain (3FK3), as well as homology models of the Swc4 SANT domain, were fitted to the 3D reconstruction of NuA4 based on EM labeling and cross-linking coupled to mass spectrometry data. The proposed locations of Eaf1 and the TINTIN module are indicated. (B) Overlay of the cryo-EM structure of Piccolo in complex with the NCP (EMD-9536) on the EM density map of full NuA4. Scale bars, 50 Å.