The Brain Bases of Phantom Auditory Phenomena ...

The Brain Bases of Phantom Auditory Phenomena: From Tinnitus to Hearing Voices Cynthia Gayle Wible, Ph.D.1

ABSTRACT

The phenomenology and neural bases of phantom auditory perceptions are reviewed. A variety of phantom auditory phenomena are discussed, from tinnitus to hearing voices. It is claimed that the phenomenology or qualia of the hallucinatory experience may correspond to how the auditory system is organized into functional regions (neural architecture) and how auditory percepts are represented at the single neuron level within this system. There may be a one-toone correspondence between the type of experience (e.g., hearing a tone versus hearing a voice) and the representational qualities or aspects of sound representation in different parts of the auditory and speech processing stream or system. The literature does not support the supposition that certain features of auditory hallucinations correspond to either neurological or psychiatric disease. Clinical aspects of auditory hallucinations also are discussed that may be of interest to clinical practitioners with patients who have auditory hallucinations. KEYWORDS: Auditory hallucinations, tinnitus, music hallucinations,

voices, schizophrenia

Learning Outcomes: As a result of this activity, the participant will be able to (1) identify brain regions involved in the generation of phantom auditory phenomena and (2) implement recommendations for patients who experience these perceptions.

P

hantom auditory phenomena or auditory hallucinations consist of auditory perception in the absence of external stimulation. Auditory

hallucinations range from hearing short simple sounds to hearing voices (speech). Voice hallucinations also range from simple short-lived

1 Department of Psychiatry, Laboratory for Neuroscience, Harvard Medical School, VA Boston Healthcare System, Brockton, Massachusetts. Address for correspondence and reprint requests: Cynthia G. Wible, Ph.D., Department of Psychiatry, Laboratory for Neuroscience, Harvard Medical School VA Boston Healthcare System, 116A, 940 Belmont Street Brockton, MA 02301 (e-mail: [email protected]).

Proceedings of the 5th International NCRAR Conference—Expanding Our Horizons: Medical Conditions and Audiology; Guest Editors, Robert L. Folmer, Ph.D., and Gabrielle H. Saunders, Ph.D. Semin Hear 2012;33:295–304. Copyright # 2012 by Thieme Medical Publishers, Inc., 333 Seventh Avenue, New York, NY 10001, USA. Tel: +1(212) 584-4662. DOI: http://dx.doi.org/10.1055/s-0032-1315728. ISSN 0734-0451.

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phenomena (hearing one’s name called) to hearing full sentences or dialogues and the feeling of a presence of a being or entity. This apparent continuum of complexity may be related to the characteristics of the brain regions that are involved in the hallucination; this will be discussed in the next section. Auditory hallucinations can arise for several reasons, but are thought to arise from central brain dysfunction or abnormal activity in the brain. Even tinnitus, a simple form of auditory hallucination, which is characterized by hearing a buzzing, ringing, or tone sound, is thought to result from abnormal brain activity and is not usually alleviated by peripheral treatments.1 So, although tinnitus is often precipitated by hearing loss or peripheral damage, the experience of hearing tinnitus sound(s) comes from abnormal activity in the brain. Tinnitus is common and has been estimated to occur in up to 20% of the population.2 Like tinnitus, musical hallucinations are most often associated with either hearing loss or neurological damage, but this is not the case with voice hallucinations.3 If voices are heard in neurologically impaired patients, they are often localized on one side and hence do not have the same experiential quality as hearing a “real” voice.4 Auditory hallucinations of voices or auditory verbal hallucinations (AVH) are the most frequent symptom of schizophrenia. Bleuler5 observed that “Almost every schizophrenic who is hospitalized hears voices, occasionally or continually.”p. 367 Silbersweig and Stern6 estimated that up to 74% of schizophrenic patients hear voices, which can be experienced as conversing with each other or commenting on ongoing behavior and are often sentences or dialogs, not just single words. Schizophrenic AVH are usually very different from the common auditory hallucination of hearing one’s name that occurs in neurologically healthy individuals.7 Schizophrenic AVH or voices are experienced as coming from a person or a presence and seem “real” as if someone is actually speaking. There is often an elaborate system of beliefs and attributions concerning the voice and its origins. There may be more than one voice and the voices can have conversations with each other. Some patients hear voices that comment on their ongoing behavior. Some have the feeling

that the voice constitutes another person inside of their body. The voices also may have a frightening or punitive tone. A first-person account of what it is like to hear voices was in a special issue of the journal Cognitive Neuropsychiatry that was dedicated to the topic of auditory hallucinations; the following is a quote from an article by Cockshutt8p. 9: For me, and I can only speak for myself, the voices are externalized. . . and real. There is no point in pretending otherwise. I could say that I understand that they are a false manifestation of my internal thoughts. The truth is that for me that is the unreal aspect of it all because by pretending to believe that the voices are unreal I am, in essence, creating a false reality. In summary, auditory hallucinations form a continuum of complexity. On one end are simple tones and noises, followed by music, which is more complex and organized. Voice hallucinations may be next in the continuum and can be categorized within the speech system as simple (hearing brief single words or unintelligible speechlike sounds) or more complex (hearing a voice or voices speaking full sentences or dialogues accompanied by a feeling that the voice has an external source or is real or corporeal). The next section of this article will describe representational aspects of the auditory speech system in the brain that may correspond to the phenomenological aspects of auditory hallucinations. THE BRAIN BASES OF PHANTOM AUDITORY PHENOMENA: TONES AND SPEECH In the previous section, a continuum of complexity in auditory hallucinations was described that can range from hearing simple tones to hearing a voice with a feeling of a source or a feeling that it comes from a real person. The later type of hallucination is most often associated with schizophrenia (hearing a voice narrative with a feeling of a source or presence). However, brain lesions can produce schizophrenia-like voice hallucinations.9 Therefore, the type of hallucination experienced may

THE BRAIN BASES OF PHANTOM AUDITORY PHENOMENA/WIBLE

depend on the brain areas involved, not the type of disease. The cortex is made up of small functional regions of neurons that respond similarly; the size and juxtaposition of these cortical maps is what I refer to as “architecture.” The adjacency or cortical location of a map can convey an abundance of information about what type of inputs it receives, the output of the computation, and the function of the region. For example, neurons within a region might all respond to faces more than other visual attributes or they might respond to visual objects and also exhibit size and shape constancy. These attributes constitute information about the neuronal representation. The size and shape of these regions varies considerably, especially beyond primary cortical regions where a columnar organization may be present. I will review evidence that the continuum of complexity in auditory hallucinations matches the representational structure and cortical architecture of progressively higher-order auditory cortical regions. This type of correspondence between hallucinations and cortical architecture and representation has been found in the visual system. Unlike language, the visual system in nonhuman primates is comparable to humans and has been mapped in great detail. Visual cortex is divided into different regions whose neurons respond primarily to color, objects, faces, and other visual categories. A one-to-one correspondence was found between the type of visual hallucination experienced in human subjects and activity in visual regions such that color hallucinations were associated with activity in visual regions representing color, face hallucinations were associated with activity in those cortical regions representing faces, and so on.10,11 Within this framework, hearing simple tones or ringing sounds would correspond to a dysfunction of more primary, tonotopically organized regions of cortex. It is well known from single-unit or single-neuron recording in animals that the primary auditory cortex is tonotopically organized into maps where neurons within a small region respond optimally to tones within a specific frequency and nearby regions respond to surrounding frequencies. Mirror symmetric tonotopic maps resembling those previously found in macaque monkeys

have now been found in the human primary auditory cortex.12 These investigators used very high-resolution functional magnetic resonance imaging (FMRI) and found mirror symmetric tonotopic maps in Heschl’s gyrus or primary auditory cortex that shared a low frequency border. Primary auditory cortex has been implicated as a generator of tinnitus in animals and human subjects. For example, the modulation of (both excitatory and inhibitory) activity in primary auditory cortex was found to be the basis of tinnitus in a recent report using an animal model whose results replicated previous work.13 If primary auditory cortex is responsible for the experience of tinnitus, then interventions that change brain activity in this region should affect tinnitus in human subjects. Human neuroimaging and magnetoencephalography studies show that the primary auditory cortex is reorganized in subjects with tinnitus. There is an expansion of the frequency representation in the auditory cortex that corresponds to the perceived tinnitus frequencies; the degree of the shift is related to the severity of the tinnitus.14,15 These findings have been used to successfully treat tinnitus in a patient with auditory nerve damage who was deaf in the left ear. FMRI was used to map out the auditory response in primary auditory cortex and to visualize the abnormally activated regions. This FMRI activity was used as a guide to place electrodes for focal extradural electrical stimulation of the primary auditory cortex. This procedure was reported to suppress tinnitus completely.15 Recently, researchers have begun to investigate hypotheses about the interactions between other regions and auditory cortex in inhibiting tinnitus, but current evidence suggests that the perception is clearly generated within the primary auditory cortex.16 Hence, abnormal activity or overactivation in the primary auditory cortex that is tonotopically organized corresponds to the experience of hearing tones or simple sounds when none are present in the environment. The next section will explore the possibility that more complex auditory hallucinations such as speech are also a result of neural overactivation and that the phenomenological aspects of the hallucinations can be linked to the neuronal representational properties and architecture of higher-order cortical

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regions that are further up in the stream of auditory processing. NEURAL BASES OF SPEECH PERCEPTION AND PRODUCTION This describes progressively higher-order auditory processing stages that form the neural basis of speech perception. Although these regions are subsequent to primary auditory cortex, the processing of information is thought to be highly recursive and interactive and does not proceed in a strictly linear fashion. For speech perception, a spectral-temporal analysis of the auditory signal is performed in primary auditory cortex, and an auditory phonological representation is activated in the middle portion of the superior temporal sulcus (STS; see Fig. 1). A more posterior region of the STS houses amodal (or multimodal) phonological representations and can be activated by written or spoken words. To initiate speech production, these representations are then translated into prearticulatory motor codes for the vocal tract in the Sylvian parietal-temporal area (Spt), a region near the temporal-parietal boundary (this summary is based on a model and synthesis from Hickok and Poeppel17,18). The Spt interacts with the middle and inferior frontal regions to produce the articulatory codes for speech production. The posterior STS and Spt are active during speech perception, production, and during subvocal rehearsal for working memory.

Phonemic representations in the STS also make contact with widespread semantic representations in the temporal lobe and other regions. Overactivation that extends beyond primary auditory cortex would be predicted to cause the perception of phonemes (perhaps in the form of incomprehensible words) or whole single-word auditory representations. Hug et al4 described an epileptic patient who exhibited a progressive auditory hallucination such that tonal tinnitus transformed into the perception of noise and then finally to incomprehensible voices, as would be predicted from the structure of the auditory speech system. Voices are usually experienced within a social context and within conversation with another person. Under naturalistic conditions, auditory voice and visual face and body gestures are experienced simultaneously. The audiovisual nature of the speech signal is reflected in an area that is in the posterior region of the STS (PSTS). A large portion of this region is dedicated to audiovisual speech processing in human subjects. This area (especially the right PSTS) functions in the recognition of voices, as opposed to recognizing verbal or semantic content.19 The PSTS is part of a system that extends upward into the inferior parietal region where intentions are formed and sent to motor regions of the brain. The PSTS and inferior parietal regions are often referred to as the “temporal-parietal occipital junction” (TPJ). This system, or collection of tightly functionally

Figure 1 Several posterior language regions that are involved in the perception and production of speech17,18. The primary auditory cortex (green), a middle portion of the superior temporal sulcus (STS; yellow) that houses auditory phonological representations. An amodal phonological region (dark blue) in the posterior STS that is involved in amodal phonological representation. The Sylvian parietal-temporal area in light blue only in the left hemisphere provides the interface between the phonological networks in the bilateral superior temporal gyrus and STS and the articulatory networks in the anterior or prefrontal language system.


coupled regions, has interesting representational and architectural properties that may correspond to characteristics present in schizophrenia-like voice hallucinations. The audiovisual signal that conveys speech also contains information about person identity (agency), as well as information about intentions and emotional state (in the form of prosody and emotional gestures). TPJ functionality reflects the fact that the voice is experienced simultaneously with social representations of persons and emotion. A dominant role of this region is in the representation of dynamic multimodal gestures (primarily sight, sound, and touch), including audiovisual speech.20 Audiovisual speech representation is adjacent to or partially overlapping in the TPJ with the neural territory responsible for prosody perception (prosody is the melodic quality of the voice that can convey both emotion and meaning), the perception of emotional expressions, the perception of eye gaze, and the perception of social attention.21–27 The coding of agency (or intention or purpose) is automatically activated or perceived along with gestures or movements and is an inherent part of the representation. Jellema et al28 recorded neuronal activity within monkey STS that combined information about reaching or grasping with activity related to attention or gaze direction. This combination of information results in a cellular representation of the intentionality of movements.28 FMRI studies in human subjects show that the PSTS is the only brain region in humans that responded differentially to intentional versus unintentional movement, leading to the conclusion that this region is a core substrate for conveying the perception of agency.29 The consequence of this coupling of gesture with a code for agency is that if the speech representations in TPJ were erroneously activated, then there would be an accompanying feeling of agency or a feeling of someone acting. Hence, overactivation of audiovisual speech gestures could cause the perception of a voice and the feeling of a presence with intentions. This could be the basis of a feeling of a source or presence that accompanies the auditory hallucination of a voice in schizophrenia-like voice hallucinations. This could be an underlying reason for the often elaborate

system of beliefs about the purpose and origin of the voice. There is direct evidence for these suppositions: Cortical stimulation of the TPJ in a nonpsychotic human subject produced a feeling of a shadowy presence, and the subject imbued this presence with certain intentions.30 In other words, the voice hallucinations feel real because they activate the part of cortex that corresponds to the perception of speaking to another person; at this cortical level of processing, the intention of the person and the audiovisual speech signal are encoded and perceived automatically and simultaneously. When audiovisual representations are activated in the PSTS, neurons provide rapid feedback input to unimodal sensory regions and especially to the auditory cortex in both animals and humans.26,27,31 This feedback is automatic and occurs without conscious attention. Hence, the overactivation or erroneous activation of audiovisual speech representations excites earlier auditory regions and could perpetuate abnormal neural activity. Another aspect of schizophrenia-like voice hallucinations is that the voices often speak in dialogues or narratives. The TPJ (bilaterally) is preferentially involved in narrative comprehension, as compared with word or even sentence comprehension.28 Therefore, this region may be used to both perceive and construct narratives. This observation might account for the fact that schizophrenic subjects are deficient in generating or building linguistic context and in understanding narrative.32 The TPJ is also selectively involved in the theory of mind or the ability to attribute and represent other’s mental states (also an important part of social communication and understanding speech and actions) as well as in selfrepresentation.21,22,25 Figure 2 shows the overlapping functionality in this region and depicts the approximate cortical regions for many of the functions discussed in this section.33–38 In fact, the TPJ may be the core region in the brain that underlies the perception of social interaction.24,33 Simply put, overactivation of this region may cause the perception of social interaction such as being in a conversation with another person or hearing a conversation. As discussed previously, the overlap between voice and dynamic person and emotion

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Figure 2 Summary figure of the overlap of functional regions in the temporal-parietal occipital junction (inferior parietal and posterior region of the superior temporal sulcus) involved in eye gaze (red); audiovisual speech (light blue); self-representation (yellow); theory of mind/agency (green); emotional perception of faces and prosody (dark blue). Rerepresentation of data respectively from Nummenmaa et al,34 Wright et al,35 Blanke and Arzy,36 Young et al,37 and Adolphs et al.38

representation may be the basis for the experience of a voice and of a presence that constitutes schizophrenia-like auditory hallucinations. There are several lines of evidence that the TPJ is involved in schizophrenia-like auditory hallucinations.26,27 First, a lesion in the TPJ, or epilepsy, can cause schizophrenia-like psychoses, including AVH.9,27 One of the most accurate but difficult ways to study AVH is the symptom capture method. In this method, the brain is imaged during the hallucination and also during periods when the hallucination is absent. One of the best symptom capture studies of schizophrenic voice hallucinations was performed using a patient whose hallucinations had a periodicity (the hallucination would last for 26 seconds with a period of silence for 26 seconds) that matched well with the requirements of FMRI imaging.39 This study showed that activity in the PSTS and middle temporal region was evident immediately before the hallucination. Also, the neural activity persisted throughout the hallucination and spread to inferior parietal and then to frontal cortex. This case study is consistent with the supposition that schizophrenia-like auditory hallucinations of voices arise from activity in the TPJ.40 Additional supporting evidence for this theory comes from reports

that transcranial magnetic stimulation applied to the TPJ has been found to alleviate schizophrenic auditory hallucinations (as well as other symptoms).41 Transcranial magnetic stimulation uses a coil to apply electromagnetic energy to the patient’s scalp/skull and has the ability to suppress or alter patterns of neural activity in brain regions beneath the coil. MUSIC HALLUCINATIONS Where in the auditory processing stream does the perception of music occur? There is relatively little research on the perception of musical hallucinations and on music perception in general, apart from other types of auditory perception. Apparently, the neural territory for music perception overlaps with speech perception in the superior temporal cortex.42,43 However, music was found to activate more dorsomedial regions, including insular and inferior parietal regions.42 A review of musical hallucinations concluded that they can result from underlying pathology in the ear or the brain.44 The findings that music activates more dorsomedial regions are consistent with a report of a patient who developed musical hallucinations after resection of a right insular glioma.45 However, additional studies are needed to


determine the underlying neural basis of music hallucinations because the findings are somewhat divergent in this area of research.46,47 CLINICAL ASPECTS AND IMPLICATIONS OF AUDITORY HALLUCINATIONS Auditory hallucinations—especially music and voice hallucinations—are thought to be underreported. Brain regions involved in producing auditory hallucinations also are involved in normal hearing, auditory attention, and working memory tasks. Therefore, all of these functions may be impaired in patients with auditory hallucinations. These factors should be considered when evaluating patients who may be experiencing hallucinations. Oliver Sacks describes several case studies of persons with musical hallucinations in his book Musicophilia,48 which provides insight into these unusual experiences, including the emotional reactions and coping strategies of patients with music hallucinations. This book is recommended reading for patients with music hallucinations, their family members, and health care providers. Auditory hallucinations can be associated with various etiologies, including brain damage, epilepsy, psychiatric disorders, or hearing loss. One report of patients referred for hearing impairment showed a high prevalence of auditory hallucinations (33%) that consisted of “humming or buzzing (35.9%), shushing (12.8%), beating or tapping (10.6%), ringing (7.7%), other individual sounds (15.4%), multiple sounds (12.6%), voices (2.5%) or music (2.5%).”49p. 444 Patients who experience music or voice hallucinations would probably benefit from a neurological or psychiatric consultation. This is especially true if the hallucinations are present for extended periods (months or longer) and are distressful or interfering with daily function. One study indicated that patients who hear auditory hallucinations, and especially voices, felt that they would benefit if they could describe the experience in more detail with health practitioners.50 Clinicians asking questions about the content and sensory aspects of the hallucination may comfort patients and may also provide valuable information about treatment strategies and referrals.

Voice hallucinations in particular can be a prominent and sometimes defining feature of schizophrenia. Schizophrenia is often accompanied by other symptoms such as hallucinations of people and delusions (e.g., people are spying on me, watching me, etc.) as well as affective inexpressiveness and attention deficits. However, voice hallucinations that comment on the patient’s actions or that consist of hearing two or more voices conversing may alone indicate schizophrenia if they are experienced for more than 2 months (see the Diagnostic and Statistical Manual of Mental Disorders for diagnostic criteria and more specific information). Because voices can sometimes command patients to harm themselves or others, clinicians should be vigilant for this type of hallucination. Strategies exist that can be used to assess the negative characteristics of unpleasant voices and to assess the potential risk resulting from hearing voices.51 Behavior management protocols (such as listening to music and other coping skills) can be taught to patients with persistent auditory hallucinations.51 SUMMARY This article reviewed evidence for understanding auditory hallucinations within a framework that is based on the functional architecture and single-neuron representational properties of the auditory processing stream. Hallucinations of short-lived sounds may correspond to the aberrant activation of more primary auditory regions. Hallucinations of voices and a feeling of a presence or that someone is speaking may correspond to the activation of audiovisual speech representations in the TPJ. In the normal brain, this region is activated when someone is actually speaking. These gesture representations automatically convey agency and intention and synchronize speech regions. The TPJ also is used to perceive and construct narratives, a feature that may account for the elaborate conversational nature of schizophrenia-like auditory voice hallucinations. Auditory hallucinations are frequently reported by patients in audiology clinics. The brain regions that were described to be involved in auditory hallucinations are also used for speech perception, speech

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production, and working memory. Some of these neural regions are used for affect perception and production (in the form of gestures and prosody) and social attention. Therefore, clinicians should be sensitive to the fact that hallucinations may interfere with these functions. Patients who experience hallucinations report a need to discuss the phenomenological aspects of the hallucinations and their meaning with caretakers. An assumption underlying the framework described in this article is that the experiences seem very real and are not under the control of the patient. The type of auditory hallucination (simple or schizophrenia-like) does not show a one-to-one correspondence to the cause (e.g., psychosis, epilepsy, hearing loss). This observation should be kept in mind for patients who are referred to other specialties (neurology, psychiatry) because the intensity or type of hallucination or distress caused by the hallucination need to be considered to identify and implement an effective treatment program. ACKNOWLEDGMENTS

This work was supported by an NIMH grant: 1 R01 MH067080–01A2 and by the Harvard Neuro-Discovery Center (formally HCNR). This work was also funded by the Biomedical Informatics Research Network (U24RR021992), National Institute of Mental Health. CONFLICT OF INTEREST STATEMENT

The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. REFERENCES 1. Folmer RL, Griest SE, Martin WH. Chronic tinnitus as phantom auditory pain. Otolaryngol Head Neck Surg 2001;124:394–400 2. Demeester K, van Wieringen A, Hendrickx JJ, et al. Prevalence of tinnitus and audiometric shape. BENT 2007;3(Suppl 7):37–49 3. David AS. The cognitive neuropsychiatry of auditory verbal hallucinations: an overview. Cogn Neuropsychiatry 2004;9:107–123

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