The mammalian inner ear separates sounds by their frequency content, and this separation underlies important properties of human hearing, including our ability to understand speech in noisy environments. Studies of genetic disorders of hearing have demonstrated a link between frequency selectivity and wave properties of the tectorial membrane (TM). To understand these wave properties better, we developed chemical manipulations that systematically and reversibly alter TM stiffness and viscosity. Using microfabricated shear probes, we show that (i) reducing pH reduces TM stiffness with little change in TM viscosity and (ii) adding PEG increases TM viscosity with little change in TM stiffness. By applying these manipulations in measurements of TM waves, we show that TM wave speed is determined primarily by stiffness at low frequencies and by viscosity at high frequencies. Both TM viscosity and stiffness affect the longitudinal spread of mechanical excitation through the TM over a broad range of frequencies. Increasing TM viscosity or decreasing stiffness reduces longitudinal spread of mechanical excitation, thereby coupling a smaller range of best frequencies and sharpening tuning. In contrast, increasing viscous loss or decreasing stiffness would tend to broaden tuning in resonance-based TM models. Thus, TM wave and resonance mechanisms are fundamentally different in the way they control frequency selectivity.cochlear mechanics | traveling waves | resonance | tectorial membrane | viscoelastic materials T he sharp frequency selectivity of auditory nerve fiber responses to sound is a hallmark of mammalian cochlear function. This remarkable signal processing originates in the mechanical stage of the cochlear signal processing chain (1-7), as evidenced by measured motions and mechanical properties of the basilar membrane (BM) (2-9) and tectorial membrane (TM) (10-24). Although the hydromechanical mechanisms underlying BM motions have been characterized based on experimental and theoretical studies, the mechanisms underlying TM motions remain unclear.The TM is an acellular matrix that overlies the hair bundles of sensory receptor cells. Based on its strategic position above the organ of Corti, conventional cochlear models (25-29) have implicated local mechanical properties (i.e., mass, stiffness) of the TM in stimulating the sensory hair bundles of hair cells and in cochlear tuning. Recent dynamic measurements of the TM, in vitro (17,(30)(31)(32)(33) and in vivo (34), suggest that the TM supports longitudinal coupling, with large spatial extents across a broad range of frequencies. This longitudinal coupling manifests in the form of propagating traveling waves that are thought to contribute to hearing mechanisms (17,21,30,(35)(36)(37)(38)(39)(40). Genetic modification studies provide further support that the spatial extent of TM waves may play a significant role in cochlear tuning (30,32). Although these measurements, models, and genetic modification studies have confirmed the importance of TM mechanical properties in ...