Review and Perspective on Brain Bases of Tinnitus.

In advancing our understanding of tinnitus, some of the more impactful contributions in the past two decades have come from human brain imaging studies, specifically the idea of both auditory and extra-auditory neural networks that mediate tinnitus. These networks subserve both the perception of tinnitus and the psychological reaction to chronic, continuous tinnitus. In this article, we review particular studies that report on the nodes and links of such neural networks and their inter-network connections. Innovative neuroimaging tools have contributed significantly to the increased understanding of anatomical and functional connections of attention, emotion-processing, and default mode networks in adults with tinnitus. We differentiate between the neural correlates of tinnitus and those of comorbid hearing loss; surprisingly, tinnitus and hearing loss when they co-occur are not necessarily additive in their impact and, in rare cases, additional tinnitus may act to mitigate the consequences of hearing loss alone on the brain. The scale of tinnitus severity also appears to have an impact on brain networks, with some of the alterations typically attributed to tinnitus reaching significance only in the case of bothersome tinnitus. As we learn more about comorbid conditions of tinnitus, such as depression, anxiety, hyperacusis, or even aging, their contributions to the network-level changes observed in tinnitus will need to be parsed out in a manner similar to what is currently being done for hearing loss or severity. Together, such studies advance our understanding of the heterogeneity of tinnitus and will lead to individualized treatment plans.