CNS Spectrums (2017), page 1 of 7. © Cambridge University Press 2017 doi:10.1017/S1092852916000870 REVIEW ARTICLE Musical hallucinations: a brief review of functional neuroimaging findings Francesco Bernardini,1* Luigi Attademo,2 Karen Blackmon,3 and Orrin Devinsky3 1 Department of Psychiatry, Hôpital Érasme, Université Libre de Bruxelles, Brussels, Belgium Department of Mental Health, ASST Papa Giovanni XXIII, Bergamo, Italy 3 Comprehensive Epilepsy Center, Department of Neurology, New York University School of Medicine, New York, New York 2 Musical hallucinations are uncommon phenomena characterized by intrusive and frequently distressful auditory musical percepts without an external source, often associated with hypoacusis, psychiatric illness, focal brain lesion, epilepsy, and intoxication/pharmacology. Their physiological basis is thought to involve diverse mechanisms, including “release” from normal sensory or inhibitory inputs as well as stimulation during seizures, or they can be produced by functional or structural disorders in diverse cortical and subcortical areas. The aim of this review is to further explore their pathophysiology, describing the functional neuroimaging findings regarding musical hallucinations. A literature search of the PubMed electronic database was conducted through to 29 December 2015. Search terms included “musical hallucinations” combined with the names of specific functional neuroimaging techniques. A total of 18 articles, all clinical case reports, providing data on 23 patients, comprised the set we reviewed. Diverse pathological processes and patient populations with musical hallucinations were included in the studies. Converging data from multiple studies suggest that the superior temporal sulcus is the most common site and that activation is the most common mechanism. Further neurobiological research is needed to clarify the pathophysiology of musical hallucinations. Received 4 May 2016; Accepted 11 November 2016 Key words: Review, functional neuroimaging, musical hallucinations. Introduction Musical hallucinations (MHs) are uncommon phenomena characterized by intrusive and frequently distressful auditory percepts without an external source1 and are described as songs, tunes, melodies, harmonics, rhythms, and/or timbres.2 Five common causes of MHs include hypoacusis, psychiatric illness, focal brain lesion, epilepsy, and intoxication/pharmacology,3 but MHs also occur in people without auditory, neurological, or psychiatric disorders.4 The physiological basis for MHs involves diverse mechanisms, including “release” from normal sensory or inhibitory inputs as well as stimulation during seizures.5 Functional or structural disorders in diverse cortical and subcortical areas can produce MHs.6 However, in some cases, neither a localized lesion nor a functional disorder * Address for correspondence: Francesco Bernardini, Department of Psychiatry, Erasme Hospital, Université Libre de Bruxelles, Route de Lennik 808, 1070 Anderlecht, Belgium. (Email: francesco.bernardini@erasme.ulb.ac.be) can be identified. To further explore the pathophysiology of MHs, we reviewed functional neuroimaging and neurochemical imaging studies. Material and Methods We searched the PubMed electronic database for all articles up to 29 December 2015. The search terms included “musical hallucinations” combined with “functional neuroimaging” or “functional magnetic resonance imaging” or “fMRI” or “magnetoencephalography” or “MEG” or “positron emission tomography” or “PET” or “singlephoton emission computed tomography” or “SPECT” or “near-infrared spectroscopy” or “NIRS.” The search included all languages. Some 11 articles were identified.6–16 We excluded two articles that were unrelated to the topic.12, 13 In addition to the PubMed search, nine other relevant clinical case reports of functional neuroimaging studies on MHs were identified based on bibliographies and our knowledge of the subject.17–25 A final set of 18 articles, all clinical case reports, comprised the set of studies reviewed. Downloaded from http:/www.cambridge.org/core. Newcastle University, on 21 Dec 2016 at 00:21:15, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S1092852916000870 2 BERNARDINI ET AL. Results Among the 18 functional neuroimaging studies published on patients with MHs, 17 were single-case reports, and 1 reported 6 cases, providing data on 23 patients. Among these 23 patients, all but one were examined with a single functional neuroimage: PET (n = 10), SPECT (n = 8), fMRI (n = 2), MEG (n = 2), and SPECT and MEG (n = 1). MHs were associated with different pathologies or conditions: (1) hearing loss or deafness without other psychiatric or neurological conditions (n = 10); (2) psychiatric disorders (n = 4) (depression = 3, schizophrenia = 1); (3) epilepsy and hearing loss (n = 2); (4) depression and hearing loss (n = 1); (5) Alzheimer’s disease (n = 1); (6) temporal ischemic stroke (n = 1); (7) depression and epilepsy with hearing loss (n = 1); (8) traumatic brain injury and progressive deafness (n = 1); (9) intracranial hemorrhage (n = 1); and (10) neuropsychiatric normality (n = 1). In three cases, the onset of MHs followed a new pharmacological treatment (donepezil, amitriptyline, gentamicin) in the presence of a preexisting neurological or psychiatric disorder. Patient characteristics, the clinical features of their MHs, and the functional neuroimaging findings of the reviewed articles are summarized in Table 1, in chronological order. Figure 1 overlays findings from each study on a Montreal Neurological Institute (MNI) template brain. Discussion Musical hallucinations involve a localized network of cortical areas, with converging data from multiple studies suggesting that the superior temporal sulcus (Figure 1) is the most common site and that activation is the most common mechanism. The superior temporal sulcus is an auditory association area that is selectively activated by music and melody (as well as speech-sound processing, such as phonological mismatch26) versus such other acoustic features as pitch27 and melody.28,29 Other cortical areas activated within the network underlying MHs include the orbitofrontal, precuneus, and basal ganglia. Activation of these areas, which are all reciprocally connected with the auditory association cortex, may occur passively via efferents from the superior temporal sulcus, actively through the same pathological process that stimulated the superior temporal sulcus, or through other mechanisms. Activation of the orbitofrontal cortex may contribute to some of the emotional features associated with MHs.30,31 The precuneus is involved in several aspects of higher-order processing of music, such as harmonic elements of melody32 and musical transformations of pitch and time,33 as well as retrieval of auditory images34 and memories.35 Basal ganglia activation may reflect timing elements of musical hallucinations, such as beat and rhythm36 but likely reflect secondary activation of these subcortical structures from temporal lobe efferents and other cortical regions activated by an MH. Diverse pathological processes and patient populations with MHs were included in the studies we reviewed. Therefore, activation of the superior temporal sulcus and related network structures may have resulted from different mechanisms, with potential possibilities including release after sensory loss, spreading cortical depression in migraine, electrical activation in epilepsy, and neurochemical anomalies with psychiatric disorders. Hearing loss is the most common disorder associated with MHs. It is likely that diminished input to the primary auditory cortex disinhibits—or “releases”—the auditory association cortex, leading to pathological activation. Why musical hallucinations are so commonly produced as opposed to simple frequencies, voices, or other auditory phenomena remains uncertain. Since music is a human universal,37 present in all cultures, its recognition and production may be a hardwired neural element activated during MHs. The limitations of this review include a somewhat small number of cases that were heterogeneous in terms of imaging techniques, timing of the functional neuroimaging studies (e.g., while patients experienced MHs or not, and time after onset of symptoms), and variability of the etiology of MHs in the reported cases. Nonetheless, a point of strength of the study is that the networks of cortical and subcortical structures that emerged from the single studies tell us that the temporofrontal cortices involved in auditory and music perception are also involved during music hallucinations. The present study reviewed the functional and chemical neuroimaging studies published on patients with MHs. From our review, we found that MHs involve a localized network of cortical areas, with the superior temporal sulcus appearing as the most common site and activation as the most common mechanism. Furthermore, our findings show that MHs are associated with mostly bilateral functional changes (60%), with a small preference to the right (25%), which is consistent with previous studies2,38 but in contrast with the largest case series of MHs to date.39 However, cross-sectional population-based studies would be useful to clarify the pathophysiology of MHs. Further prospective studies following patients with MHs compared to age-matched controls with a standardized neuroimaging assessment would be helpful to better understand the phenomenon of MHs. Furthermore, given the lack of a method to experimentally manipulate the intensity of hallucinations, the use of a residual inhibition paradigm (i.e., a transient suppression of a phantom percept after the offset of a masking stimulus) could be empirically tested in future studies of subjects with MHs to better understand this phenomenon.6 Downloaded from http:/www.cambridge.org/core. Newcastle University, on 21 Dec 2016 at 00:21:15, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S1092852916000870 Study Age (years), gender Prior disorder Clinical features of MHs Functional neuroimaging performed exam(s) Findings Erkwoh et al., 19937 55, M Depression Unknown PET performed during MHs Kasai et al., 19998 88, F No psychiatric, neurological conditions. No hearing impairment Onset abrupt. Hears various tunes of familiar favorites Magnetometry and SPECT both performed during MHs and in their absence Terao & Matsunaga, 199920 75, F Moderate bilateral hearing loss Bilaterally increased metabolism in the superior temporal cortex, parasagittal occipital cortex, thalamus, and bilateral basal ganglia Magnetometry: specific functional changes in the right auditory association cortex during MHs SPECT: Adjusted RCBF was greater in the right superior temporal and the right inferior frontal gyri during MHs Slight hypoperfusion in the bilateral thalamus and basal ganglia Griffiths, 20009 73, M Griffiths, 20009 71, M Griffiths, 20009 78, F Griffiths, 20009 58, F Griffiths, 20009 65, M Griffiths, 20009 82, F Abrupt onset of continuous MHs of old Japanese SPECT songs, associated with palinacousis 5 years symptomatic hearing loss Abrupt onset of continuous MHs in the form of 12 PET scans performed at Group analysis of the PET results was carried out for the four subjects showing multiple persons singing familiar melodies different times; for each scan, similar experiences during scanning, age, handedness, and musicality. There was with indistinguish-able lyrics the patient was asked to rate no correlation between MH strength and activation of Heschl’s gyrus on either side the severity of MHs during the in the group analysis. There was significant activation with MHs in both planum scan temporale, right basal ganglia, and right frontal operculum, the posterior temporal lobes (mainly right), both cerebellar lobes, the left deep Sylvian cortex, and the left frontal lobe 40 years symptomatic hearing loss Abrupt onset of continuous MHs of light opera pieces and popular songs 40 years symptomatic hearing loss Abrupt onset of almost continuous MHs of organ or piano music sometimes accompanied by singers; if accompanied by singers, the lyrics were distinguishable 23 years symptomatic hearing loss Gradual onset of almost continuous MHs of individual notes with the quality of a buzzy pitch 40 years progressive deafness Continuous MHs of 3 or 4 males singing familiar This subject was excluded from the analysis because of a lack of variability in MH songs with musical instruments; onset severity followed shortly after a head injury, and was accompanied by hearing a localized noise behind his head and experiencing a hot feeling that rose up the back of his head on two occasions only 15 years of progressive deafness Continuous MHs of one or more singers and This subject was excluded from the analysis because of markedly abnormal brain accom-panying piano or band; onset structure (large arachnoid cyst in the right occipital lobe) coincident with an episode of visual loss, disorientation, perplexion, slurring of speech, unsteadiness of body; this is likely to have been a posterior circulation vascular event; she also experienced environmental sound illusions and verbal hallucinations FUNCTIONAL NEUROIMAGING IN MUSICAL HALLUCINATIONS 3 Downloaded from http:/www.cambridge.org/core. Newcastle University, on 21 Dec 2016 at 00:21:15, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S1092852916000870 TABLE 1. Characteristics of 23 patients with MHs and functional neuroimaging findings Study Age (years), gender Prior disorder Clinical features of MHs Tanriverdi et al., 200110 38, F Izumi et al., 200217 51, M Past history of grand mal seizures; Abrupt hearing loss and MHs in the form of untreated depression lasting popular songs after an i.m. injection of over a year gentamicin Bilateral hearing impairment Verbal and MHs (popular songs) Shinosaki et al., 200318 78, F Kopeček et al., 200519 60, F Mori et al., 200611 73, F Umene et al., 200821 83, M Shoyama et al., 201014 52, F Cosentino et al., 201022 63, M Functional neuroimaging performed exam(s) Findings SPECT (performed after the Hypoperfusion of the left lateral temporal lobe initiation of moclobemide therapy) Three SPECTs were performed with Increased RCBF in both lower frontal areas and both basal ganglia during MHs the patient in different conditions (baseline without hallucinations, MHs, and verbal hallucinations) Progressive hearing impairment; Abrupt onset of almost continuous MHs in the Eight MEGs performed in the Desynchronization in the right auditory cortex, including Heschl’s gyrus planum depression form of familiar melodies with accompanying patient and in four healthy temporale and supramarginal gyrus, during MHs musical instruments; at the time of the MEG controls. study, the MHs were replaced by simple rhythmic sounds 18 Epilepsy, anteromesio-basal MHs started at age 56 and later developed FDG PET Right anterior temporal hypometabolism (reflecting prior resection) and increased resection of the right temporal obsessive-compulsive disorder metabolism in the orbitofrontal cortex, middle/inferior frontal gyrus bilaterally, lobe and hypoacusis and left nucleus putamen; auditory cortices were normal AD MHs of familiar songs began after starting SPECT performed during MHs and Compared to nine controls, RCBF was significantly increased in the left superior donepezil (5 mg/day) in nine sex-, age-, and temporal and left angular gyri cognitive function-matched AD patients without delusions and hallucinations Major depressive disorder with MHs of old popular Japanese songs at the SPECT Areas of hypoperfusion in the bilateral temporal lobe and basal ganglia psychotic features beginning of his depressive state Persistent lower abdominal pain MHs of old familiar nursery rhymes with accom- SPECT performed before and Following treatment, as compared to the initial scan, RCBF was decreased in the and depression after lithotripsy panying musical instruments or only the 43 days after starting lenticular nucleus, thalamus, and hippocampus, and increased throughout most carbamazepine treatment of the neocortical and cerebellum over both hemispheres; the increase was for a right ureteral calculus melodies (sung by an uniden-tified but familiar voice) often occurred during the quiet maximal in the left angular region, while the maximum decrease was in the right thalamus of the night developed after a treatment with amitriptyline (85 mg/day); MHs decreased gradually and completely disappeared at 30 days after beginning carbamazepine treatment Cranial trauma and right temporal Continuous MHs in the form of popular Italian PET A brain MRI performed after the trauma revealed an area of increased signal injury in a patient with songs appeared a few days after a road intensity on T2-weighted images indicating a contusion of the right temporal pole progressive bilateral deafness accident, causing cranial trauma and right PET scans of the brain showed a hypoactive area corresponding to the temporal that started at least 20 years temporal injury; the sound volume of MHs was lesion evidenced on the MRI scan, while increased focal FDG uptake was detected earlier initially low and then became progressively in the right posterior temporal lobe louder; MHs changed in severity over the course of the day 4 BERNARDINI ET AL. Downloaded from http:/www.cambridge.org/core. Newcastle University, on 21 Dec 2016 at 00:21:15, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S1092852916000870 TABLE 1. Continued 35, M Schizo-phrenia Auditory hallucinations, musical in content and obsessive in form Calabrò et al., 201216 82, F Right temporal infarction Complex MHs of popular Italian songs increasingly in intensity during the time Vitorovic & Biller, 201323 60, F Giermanski et al., 201324 30, M Bilateral sensory-neural hearing loss for a few years Intracranial hemorrhage secondary to pineal choriocarcinoma Kumar et al., 20146 66, F 20 years hearing loss Suddenly developed MHs in the form of recognizable and unrecognizable songs Childhood onset of MHs two years after SPECT intracranial hemorrhage in the form of repeating loops of familiar song frag-ments; the onset of MHs could be spon-taneous or triggered by a true auditory musical perception MHs of piano melodies without vocals MEG during different states Futamura et al., 201425 83, M Three years symptomatic Suddenly developed MHs in the form of children, SPECT sensorineural bilateral hearing folk, military songs and the Japanese national loss; epilepsy anthem fMRI (while the patient experienced MHs) Increased activation of inferior and middle frontal gyri bilaterally, left dorsolateral prefrontal cortex, right orbitofrontal cortex, and right middle temporal gyrus. In particular, an increased activation of the right auditory associated cortex and striatal regions, primarily the left caudate head fMRI performed in the patient and Activation involving the primary auditory cortex and temporal associative areas in five normal controls bilaterally in the patient and in five normal controls, a significant increased activation in temporal planum, mostly of the right temporal cortex (in the ischemic area) in the patient PET Unremarkable Unremarkable Four left brain regions showed increased oscillatory activity during higher MH intensity compared to low MH intensity; significant power changes, after wholebrain correction, in theta/alpha, beta, and gamma bands, but not the delta or high gamma bands; increased gamma band maximal in the left anterior superior temporal gyrus; increased beta band maximal in left motor, posterior cingulate, precuneus and retrosplenial cortices; in a combined theta and alpha band, power increased maximal in the left lateral orbitofrontal cortex Late phase brain SPECT showed decreased accumulation in the right temporal lobe compared to the early phase AD = Alzheimer’s disease; 18FDG = 18-fluoro-D-glucose; fMRI = functional magnetic resonance imaging; MEG = magnetoencephalography; MHs = musical hallucinations; PET = positron emission tomography; RCBF = regional cerebral blood flow; SPECT = single-photon emission computed tomography. FUNCTIONAL NEUROIMAGING IN MUSICAL HALLUCINATIONS 5 Downloaded from http:/www.cambridge.org/core. Newcastle University, on 21 Dec 2016 at 00:21:15, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S1092852916000870 Bleich-Cohen et al., 201115 6 BERNARDINI ET AL. FIGURE 1. Schematic display of overlapping results from all studies reviewed. Binary masks were created for each region that showed increased activation in association with musical hallucinations from each study. Masks were created in standard MNI 1-mm template space using the Harvard–Oxford Cortical and Subcortical Structural Atlases and Juelich Histological Atlas, as available from the FMRIB Software Library (http://fsl.fmrib.ox.ac.uk/fsl/fslwiki/). Binary masks were summed to create an overlap map that depicts the number of studies showing positive findings at any given voxel. Red represents at least one report of positive activation at that voxel, whereas yellow represents positive findings in at least six studies. Disclosures Francesco Bernardini, Karen Blackmon, Luigi Attademo, and Orrin Devinsky hereby state that they have no conflicts of interest to disclose. R E F E RE N C E S : 1. Blom JD, Coebergh JA, Lauw R, Sommer IE. Musical hallucinations treated with acetylcholinesterase inhibitors. Front Psychol. 2015; 6: 46. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4387860/pdf/ fpsyt-06-00046.pdf. 2. Berrios GE. Musical hallucinations: a historical and clinical study. Br J Psychiatry. 1990; 156: 188–194. 3. Evers S, Ellger T. The clinical spectrum of musical hallucinations. J Neurol Sci. 2004; 227(1): 55–65. 4. Zabalza-Estevez RJ. Alucinaciones musicales: la música perpetua [Musical hallucinations: perpetual music] [in Spanish]. Rev Neurol. 2014; 58: 207–212. http://www.revneurol.com/sec/resumen.php? or=pubmed&id=2013443. 5. Coebergh JA, Lauw RF, Bots R, Sommer IE, Blom JD. Musical hallucinations: review of treatment effects. Front Psychol. 2015; 6: 814. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4468361/ pdf/fpsyg-06-00814.pdf. 6. Kumar S, Sedley W, Barnes GR, Teki S, Friston KJ, Griffiths TD. A brain basis for musical hallucinations. Cortex. 2014; 52: 86–97. Epub ahead of print Dec 17, 2013. https://www.ncbi.nlm.nih.gov/ pmc/articles/PMC3969291/pdf/main.pdf. 7. Erkwoh R, Ebel H, Kachel F, et al. 18FDG–PET and electroencephalographic findings in a patient suffering from musical hallucinations. Nuklearmedizin. 1993; 32(3): 159–163. 8. Kasai K, Asada T, Yumoto M, Takeya J, Matsuda H. Evidence for functional abnormality in the right auditory cortex during musical hallucinations. Lancet. 1999; 354(9191): 1703–1704. 9. Griffiths TD. Musical hallucinosis in acquired deafness: phenomenology and brain substrate. Brain. 2000; 123(Pt 10): 2065–2076. http://brain.oxfordjournals.org/content/123/10/ 2065.long. 10. Tanriverdi N, Sayilgan MA, Ozçürümez G. Musical hallucinations associated with abruptly developed bilateral loss of hearing. Acta Psychiatr Scand. 2001; 103(2): 153–155. 11. Mori T, Ikeda M, Fukuhara R, et al. Regional cerebral blood flow change in a case of Alzheimer’s disease with musical hallucinations. Eur Arch Psychiatry Clin Neurosci. 2006; 256(4): 236–239. Epub ahead of print Nov 29, 2005. 12. Plewnia C, Bischof F, Reimold M. Suppression of verbal hallucinations and changes in regional cerebral blood flow after intravenous lidocaine: a case report. Prog Neuropsychopharmacol Biol Psychiatry. 2007; 31(1): 301–303. 13. Arias Gómez M. Music and neurology [in Spanish]. Neurologia. 2007; 22(1): 39–45. 14. Shoyama M, Ukai S, Kitabata Y, et al. Evaluation of regional cerebral blood flow in a patient with musical hallucinations. Neurocase. 2010; 16(1): 1–6. 15. Bleich-Cohen M, Hendler T, Pashinian A, Faragian S, Poyurovsky M. Obsessive musical hallucinations in a schizophrenia patient: psychopathological and fMRI characteristics. CNS Spectr. 2011; 16(7): 153–156. 16. Calabrò RS, Baglieri A, Ferlazzo E, Passari S, Marino S, Bramanti P. Neurofunctional assessment in a stroke patient with musical hallucinations. Neurocase. 2012; 18(6): 514–520. Epub ahead of print Jan 6. 17. Izumi Y, Terao T, Ishino Y, Nakamura J. Differences in regional cerebral blood flow during musical and verbal hallucinations. Psychiatry Res. 2002; 116(1–2): 119–123. 18. Shinosaki K, Yamamoto M, Ukai S, et al. Desynchronization in the right auditory cortex during musical hallucinations: an MEG study. Psychogeriatrics. 2003; 3(2): 88–92. http://onlinelibrary.wiley.com/ doi/10.1046/j.1479-8301.2003.00009.x/pdf. 19. Kopeček M, Brunovský M, Bareš M, et al. Regional cerebral metabolic abnormalities in individual patients with nonquantitative 18 FDG PET and qEEG (LORETA). Psychiatrie. 2005; 9(Suppl 3): 56–63. http://www.academia.edu/17947148/Regional_cerebral_ metabolic_abnormalities_in_individual_patients_with_ non-quantitative_18FDG_PET_and_qEEG_LORETA_. 20. Terao T, Matsunaga K. Musical hallucinations and palinacousis. Psychopathology. 1999; 32(2): 57–59. Downloaded from http:/www.cambridge.org/core. Newcastle University, on 21 Dec 2016 at 00:21:15, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S1092852916000870 FUNCTIONAL NEUROIMAGING IN MUSICAL HALLUCINATIONS 21. Umene W, Yoshimura R, Hori H, Nakamura J. A case of vascular depression associated with musical hallucinations successfully treated with paroxetine and a low dose of risperidone. Psychogeriatrics. 2008; 8: 38–41 https://www.researchgate.net/ publication/230006414_A_case_of_vascular_depression_ associated_with_musical_hallucinations_successfully_treated_ with_paroxetine_and_a_low_dose_of_risperidone. 22. Cosentino G, Giglia G, Palermo A, et al. A case of post-traumatic complex auditory hallucinosis treated with rTMS. Neurocase. 2010; 16(3): 267–272. Epub ahead of print Jan 25. 23. Vitorovic D, Biller J. Musical hallucinations and forgotten tunes: case report and brief literature review. Front Neurol. 2013; 4: 109. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3737457/pdf/ fneur-04-00109.pdf. 24. Giermanski J, Bastiampillai T, Gupta A. “I’ve got an iPod in my head.” Aust N Z J Psychiatry. 2013; 47(10): 963–964. Epub ahead of print Mar 14. 25. Futamura A, Katoh H, Kawamura M. Successful treatment with anti-epileptic-drug of an 83-year-old man with musical hallucinosis [in Japanese]. Brain Nerve. 2014; 66(5): 599–603. 26. Scharinger M, Domahs U, Klein E, Domahs F. Mental representations of vowel features asymmetrically modulate activity in superior temporal sulcus. Brain Lang. 2016; 163: 42–49. Epub ahead of print Sep 24. 27. Norman-Haignere S, Kanwisher NG, McDermott JH. Distinct cortical pathways for music and speech revealed by hypothesis-free voxel decomposition. Neuron. 2015; 88(6): 1281–1296. 28. Griffiths T, Buchel C, Frackowiak R, Patterson RD. Analysis of temporal structure by the human brain. Nat Neurosci. 1998; 1(5): 422–427. 29. Patterson RD, Uppenkamp S, Johnsrude IS, Griffiths TD. The processing of temporal pitch and melody information in auditory cortex. Neuron. 2002; 36: 767–776. http://www.cell.com/ neuron/fulltext/S0896-6273(02)01060-7. 7 30. Blood AJ, Zatorre RJ, Bermudez P, Evans AC. Emotional responses to pleasant and unpleasant music correlate with activity in paralimbic brain regions. Nat Neurosci. 1999; 2: 382–387. 31. Joos K, Vanneste S, De Ridder D. Disentangling depression and distress networks in the tinnitus brain. PLoS One. 2012; 7(7): e40544. Epub ahead of print Jul 12. https://www.ncbi.nlm.nih.gov/ pmc/articles/PMC3395649/pdf/pone.0040544.pdf. 32. Spada D, Verga L, Iadanza A, Tettamanti M, Perani D. The auditory scene: an fMRI study on melody and accompaniment in professional pianists. Neuroimage. 2014; 102(Pt 2): 764–775. Epub ahead of print Aug 28. 33. Foster NE, Halpern AR, Zatorre RJ. Common parietal activation in musical mental transformations across pitch and time. Neuroimage. 2013; 75: 27–35. Epub ahead of print Mar 5. 34. Yoo SS, Lee CU, Choi BG. Human brain mapping of auditory imagery: event related functional MRI study. NeuroReport. 2001; 12(14): 3045–3049. 35. Huijbers W, Vannini P, Sperling RA, Pennartz CMA, Cabeza R, Daselaar SM. Explaining the encoding/retrieval flip: memoryrelated deactivations and activations in the posteromedial cortex. Neuropsychologia. 2012; 50(14): 3764–3774. https://www.ncbi. nlm.nih.gov/pmc/articles/PMC3811140/pdf/nihms407956.pdf. 36. Merchant H, Grahn J, Trainor L, Rohrmeier M, Fitch WT. Finding the beat: a neural perspective across humans and non-human primates. Philos Trans R Soc Lond B Biol Sci. 2015; 370(1664): 20140093. https://www.ncbi.nlm.nih.gov/pmc/articles/ PMC4321134/pdf/rstb20140093.pdf. 37. Brown D. Human universals. New York: McGraw-Hill; 1991. 38. Keshavan MS, David AS, Steingard S, Lishman W. Musical hallucinations: a review and synthesis. Cogn Behav Neurol. 1992; 5: 211–223. 39. Golden EC, Josephs KA. Minds on replay: musical hallucinations and their relationship to neurological disease. Brain. 2015; 138(Pt 12): 3793–3802. Epub ahead of print Oct 7. http://brain.oxfordjournals.org/ content/138/12/3793.long. Downloaded from http:/www.cambridge.org/core. Newcastle University, on 21 Dec 2016 at 00:21:15, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S1092852916000870