The Mismatch Negativity (MMN) – a Unique Window to Disturbed Central Auditory Processing in Ageing and Different Clinical Conditions ⇑ R

Clinical Neurophysiology 123 (2012) 424–458

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Clinical Neurophysiology

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Invited Review The mismatch negativity (MMN) – A unique window to disturbed central auditory processing in ageing and different clinical conditions ⇑ R. Näätänen a,b,c, , T. Kujala c,d, C. Escera e,f, T. Baldeweg g,h, K. Kreegipuu a, S. Carlson i,j,k, C. Ponton l a Institute of Psychology, University of Tartu, Tartu, Estonia b Center of Integrative Neuroscience (CFIN), University of Aarhus, Aarhus, Denmark c Cognitive Brain Research Unit, Cognitive Science, Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland d Cicero Learning, University of Helsinki, Helsinki, Finland e Institute for Brain, Cognition and Behavior (IR3C), University of Barcelona, Barcelona, Catalonia, Spain f Cognitive Neuroscience Research Group, Department of Psychiatry and Clinical Psychobiology, University of Barcelona, Barcelona, Catalonia, Spain g Institute of Child Health, University College London, London, UK h Great Ormond Street Hospital for Sick Children NHS Trust, London, UK i Neuroscience Unit, Institute of Biomedicine/Physiology, University of Helsinki, Helsinki, Finland j Brain Research Unit, Low Temperature Laboratory, Aalto University School of Science and Technology, Espoo, Finland k Medical School, University of Tampere, Tampere, Finland l Compumedics NeuroScan, Santa Fe, NM, USA article info highlights

Article history: The mismatch negativity (MMN) indexes different types of central auditory abnormalities in different Accepted 20 September 2011 neuropsychiatric, neurological, and neurodevelopmental disorders. Available online 13 December 2011 The diminished amplitude/prolonged peak latency observed in patients usually indexes decreased auditory discrimination. Keywords: An MMN deficit may also index cognitive and functional decline shared by different disorders irrespec- The mismatch negativity (MMN) tive of their specific aetiology and symptomatology. Central auditory processing MMN deficits index deficient N-methyl-D-aspartate (NMDA) receptor function affecting memory-trace Event-related potentials (ERP) Magnetoencephalography formation and hence cognition in different disorders. Auditory discrimination Neuropsychiatric disorders abstract Neurological disorders Neurodevelopmental disorders The NMDA-receptor system In this article, we review clinical research using the mismatch negativity (MMN), a change-detection response of the brain elicited even in the absence of attention or behavioural task. In these studies, the MMN was usually elicited by employing occasional frequency, duration or speech-sound changes in repetitive background stimulation while the patient was reading or watching videos. It was found that in a large number of different neuropsychiatric, neurological and neurodevelopmental disorders, as well as in normal ageing, the MMN amplitude was attenuated and peak latency prolonged. Besides indexing decreased discrimination accuracy, these effects may also reflect, depending on the specific stimulus paradigm used, decreased sensory-memory duration, abnormal perception or attention control or, most importantly, cognitive decline. In fact, MMN deficiency appears to index cognitive decline irrespective of the specific symptomatologies and aetiologies of the different disorders involved. Ó 2011 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.

Contents

1. Introduction ...... 425 2. The MMN as an index of decreased auditory discrimination accuracy (Table 1)...... 431 3. The MMN as an index of shortened sensory-memory duration (Table 2)...... 433

⇑ Corresponding author at: Cognitive Brain Research Unit, Cognitive Science, Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland. Tel.: +358 40 501 5740; fax: +358 9 191 29450. E-mail address: risto.naatanen@helsinki.ﬁ (R. Näätänen).

1388-2457/$36.00 Ó 2011 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved. doi:10.1016/j.clinph.2011.09.020 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 425

4. The MMN as an index of abnormal auditory perception ...... 434 5. The MMN as an index of increased backward masking (Table 3)...... 434 6. The MMN as an index of abnormal involuntary attention switching: either too weak (Table 4) or too strong (Table 5)...... 434 7. The MMN attenuation caused by cerebral grey-matter loss or other structural change (Table 6)...... 435 8. The MMN as an index of pathological brain excitation/excitability (Table 7)...... 437 9. The MMN as an index of cognitive and functional deterioration (Table 8)...... 437 10. The MMN as an index of the level of consciousness (Table 9)...... 438 11. The MMN as an index of the progression/severity of the illness (Table 10) ...... 438 12. The MMN in predicting illness course/drug response (prognosis) (Table 11) ...... 439 13. The MMN as an index of genetic disposition to different pathologies (Table 12)...... 440 14. The MMN as an index of central auditory processing improvement or recovery (Table 13) ...... 441 15. Concluding discussion ...... 443 Acknowledgements ...... 444 References ...... 444

1. Introduction tion (or regularity-violation) response, the mismatch negativity (MMN; Näätänen et al., 1978; Näätänen and Michie, 1979) and Interestingly, central auditory processing is affected in a large its magnetoencephalographic (MEG) equivalent, the MMNm (Hari number of different clinical conditions with very different aetiolo- et al., 1984; Sams et al., 1985b). The MMN can be reliably (Pekko- gies and symptoms such as schizophrenia, dyslexia, stroke, speciﬁc nen et al., 1995a,b; Escera and Grau, 1996; Tervaniemi et al., 1999; language impairment (SLI), multiple sclerosis (MS), amyotrophic Escera et al., 2000b) recorded even in the absence of the subject or lateral sclerosis (ALS), epilepsy and autism, suggesting some patient’s attention or behavioural task, e.g., in sleeping infants shared aetiology in these different abnormalities (see also Gottes- (Ruusuvirta et al., 2009), stroke patients even at a very early post man and Gould, 2003; Bishop, 2009; Dolan et al., 1993; Hugdahl stroke-onset period (Ilvonen et al., 2001, 2004) and in comatose and Calhoun, 2010; Reite et al., 2009). These effects on central (Kane et al., 1993, 1996; Fischer et al., 1999; Fischer and Luauté, auditory processing in different clinical conditions and in ageing 2005) and persistent-vegetative-state (PVS) patients (Wijnen can now be objectively evaluated, and hence their degree of simi- et al., 2007). larity determined, by using the electrophysiological change-detec-

Fig. 1. Grand average difference waveforms (9 subjects) for changes in duration, frequency, and perceived sound-source location for six different magnitudes of deviance at electrode Fz and referenced to the mean of the two mastoid electrodes (upper panel). The same waveforms referenced to the nose electrode and shown from Fz and the right mastoid (RM); lower panel. Sound onset is always at 0 ms (From Pakarinen et al., 2007). 426 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458

The auditory MMN response is mainly generated by a change- modality. By contrast, the sMMN appears to have, similar to the detection process occurring bilaterally in auditory cortices, in auditory MMN, sensory-specific (somatosensory cortex) and fron- which the current auditory input is found to differ from the repre- tal subcomponents (Restuccia et al., 2009; Spackman et al., sentation of the preceding auditory events, including regularities 2010), consistent with its fronto-central, contralaterally predomi- governing consecutive stimulus events (for reviews, see Näätänen nant scalp distribution (Wei et al., 2002). Currently, it is not clear et al., 2001, 2007, 2010, 2011b; Duncan et al., 2009; Winkler, whether the auditory and somatosensory modalities share the 2007). For an illustration, see Fig. 1. In subjects performing some frontal activation (generating the frontal MMN component) proba- primary, e.g., visual, task, this preconscious auditory-cortex bly serving attention switch to stimulus change (Näätänen, 1992; change-detection process reaches conscious perception by activat- Näätänen et al., 2002; Schröger, 1997). In addition, there is also ing, with a brief delay (Rinne et al., 2000; see also Rinne et al., 2005, an oMMN, with a long peak latency (about 500–600 ms) and parie- 2006), a right frontal-cortex process which, in turn, initiates further tally predominant scalp distribution (Krauel et al., 1999). cerebral processes which may lead to conscious change detection The MMN can also be recorded in different animals [in monkey (Näätänen, 1990, 1992; Näätänen et al., 2011b; Alain et al., 1998; (Javitt et al., 1992); cat (Csépe et al., 1987, 1988, 1989; Pincze et al., Berti and Schröger, 2001; Deouell et al., 2007; Giard et al., 1990; 2001, 2002); rabbit (Astikainen et al., 2001); rat (Astikainen et al., Schröger, 1996; Escera et al., 2000a). Hence, the MMN is composed 2006; Ruusuvirta et al., 1998, 2007; Roger et al., 2009; Tikhonravov of overlapping contributions from auditory- and frontal-cortex pro- et al., 2008, 2010); guinea pig (Kraus et al., 1994); and mouse cesses (the supratemporal and frontal MMN subcomponents, (Umbricht et al., 2005; Ehrlichman et al., 2008, 2009)]. respectively) (Giard et al., 1990). These two subcomponents of the The MMN enables one to determine discrimination accuracy, MMN can be inferred from the data presented in Fig. 2 showing usually with a good correspondence with behavioural discrimina- how ethanol selectively attenuates the MMN amplitude recorded tion (Sams et al., 1985a; Amenedo and Escera, 2000; Gottselig over the frontal cortex, whereas the polarity-reversed MMN re- et al., 2004; Lang et al., 1990; Näätänen et al., 1993, 2007; Leitman corded at the mastoids (with a nose reference) is unaffected. This et al., 2010; Kujala and Näätänen, 2010; Tremblay et al., 1998), sep- data pattern suggests that the frontally recorded MMN is composed arately for each auditory dimension (such as frequency, intensity of contributions from both the auditory and frontal cortices, and duration) as well as for the different speech sounds (Kraus whereas the mastoid ‘MMN’ gets a contribution from the audi- et al., 1996), with the MMN vanishing at about the behavioural dis- tory-cortex MMN generator only. Intracranial recordings in humans crimination threshold (Sams et al., 1985a; Winkler and Näätänen, also support both auditory- and frontal-cortex MMN generators 1994; Winkler et al., 1993). Therefore, it permits one to form objec- (Halgren et al., 1995a,b, 1998; Baudena et al., 1995; Kropotov tive deterioration profiles covering all important auditory dimen- et al., 1995, 2000; Liasis et al., 1999, 2000a,b; Rosburg et al., 2005, sions in different patient groups (Näätänen et al., 2004; Kujala 2007). et al., 2005a, 2006, 2007, 2010; Pakarinen et al., 2007, 2009, 2010). The MMNm, in turn, selectively reflects the temporal-lobe com- In addition, these MMN-based objective tests can be extended to ponent of the MMN because the MEG is insensitive to the radially all aspects of short-term auditory sensory memory, too, such as its oriented frontal generators (Hämäläinen et al., 1993) of the MMN. Therefore, the MMNm is particularly well suited for detecting central auditory processing deficits specific for the temporal lobes. Consequently, the combined use of the electroencephalography (EEG) and MEG recordings helps one to separately determine to what extent the temporal and frontal MMN components are impaired (Hämäläinen et al., 1993; Näätänen, 1992). Further, these two components have their functional magnetic resonance imaging (fMRI) (Molholm et al., 2005; Celsis et al., 1999; Opitz et al., 2002; Schall et al., 2003), positron emission tomography (PET) (Tervaniemi et al., 2000; Dittmann-Balcar et al., 2001; Müller et al., 2002) and optical-imaging (OI) (Rinne et al., 1999; Tse and Penney, 2008) equivalents as well. Moreover, there is event-related potential (ERP; Paavilainen et al., 1991; Doeller et al., 2003; Frodl- Bauch et al., 1997; Escera et al., 2002), MEG (Levänen et al., 1993, 1996; Rosburg, 2003) and fMRI evidence (Molholm et al., 2005) for the attribute-specific organisation of the supratemporal MMN generator. Moreover, the MMN has its equivalents in other sensory modalities, too [the visual MMN; vMMN (Alho et al., 1992; Amenedo et al., 2007; Astikainen and Hietanen, 2009; Tales et al., 1999, 2002a,b, 2008; Tales and Butler, 2006; Maekawa et al., 2005, 2009; Berti and Schröger, 2004, 2006; Stagg et al., 2004; Stefanics et al., 2011; Heslenfeld, 2003; Pazo-Alvarez et al., 2003, 2004; Fro- yen et al., 2010; Iijima et al., 1996; Kenemans et al., 2003; Kimura et al., 2009; Czigler, 2007; Czigler and Csibra, 1990, 1992; Czigler et al., 2002, 2004, 2006a,b; Czigler and Pató, 2009; Nordby et al., 1996; Wei et al., 2002; Woods et al., 1992); the somatosensory MMN; sMMN (Kekoni et al., 1997; Shinozaki et al., 1998; Akatsuka et al., 2005; Kida et al., 2001; Spackman et al., 2007, 2010; Astikai- Fig. 2. The differential effects of ethanol on MMN at the frontal (Fz) and left and nen et al., 2001; Näätänen, 2009); and the olfactory MMN; oMMN right mastoid (LM and RM, respectively) electrode positions. MMN was reduced at the frontal electrode but the polarity-inverted MMN recorded at the mastoids, (Krauel et al., 1999; Pause and Krauel, 2000)]. The vMMN has a suggesting the supratemporal MMN subcomponent, was not affected by ethanol parieto-occipital scalp distribution but there is no convincing evi- (0.55 g/kg) whereas the frontal subcomponent was affected by ethanol. Adapted dence for a frontal component analogous to that in the auditory from Jääskeläinen et al. (1996b). Table 1 The auditory MMN (MMNm) provides an objective index of affected auditory discrimination in:

– ADHD (Alexandrov et al., 2003; Barry et al., 2003; Gumenyuk et al., 2005; Huttunen et al., 2007; Huttunen-Scott et al., 2008; Kemner et al., 1996; Kilpeläinen et al., 1999; Oades et al., 1996; Rothenberger, 1995; Rothenberger et al., 2000; Satterﬁeld et al., 1988; Sawada et al., 2010; Wild-Wall et al., 2005; Winsberg et al., 1993) – Ageing (Alain and Woods, 1999; Alain et al., 2004; Amenedo and Diaz, 1998; Bellis et al., 2000; Bertoli et al., 2005; Cooper et al., 2006; Czigler et al., 1992; Fabiani et al., 2006; Gaeta et al., 1998, 1999, 2001, 2002; Gunter et al., 1996; Horváth et al., 2007; Iijima et al., 1996; Ikeda et al., 2004; Jääskeläinen et al., 1999a; Karayanidis et al., 1995; Kiang et al., 2007, 2009; Kisley et al., 2004a,b, 2005; Kok, 2000; Mager et al., 2005; Osawa et al., 1996; Pekkonen, 2000; Pekkonen et al., 1995a, 1996a, 2005; Schiff et al., 2008; Schroeder et al., 1995; Verleger et al., 1991; Woods, 1992; Woods and Clayworth, 1986) – Alcohol intoxication, acute (Ahveninen et al., 2000a; Jääskeläinen et al., 1995, 1996a,b, 1998, 1999a,b; Kähkönen et al., 2001, 2004) Alcoholism (Ahveninen et al., 2000a,b; Fein et al., 2004; Grau et al., 2001; Holguin et al., 1998; Kathmann et al., 1995; Marco-Pallares et al., 2007; Pekkonen et al., 1998; Polo et al., 2003; Realmuto et al., 1993; Rodriguez et al., 1998; van der Stelt and Belger, 2007; Zhang et al., 2001) – Alzheimer’s disease and dementia (Engeland et al., 2002; Gaeta et al., 1999; Katada et al., 2004; Kazmerski et al., 1997; Missioner et al., 1999; Pekkonen, 2000; Pekkonen et al., 1994, 1996b, 1999, 2001a,b; Riekkinen et al., 1997; Schroeder et al., 1995; Yokoyama et al., 1995) – Amusia(Congenital) (Moreau et al., 2009; Peretz et al., 2009) – Amusia (Caused by Stroke) (Kohlmetz et al., 2001; Särkämö et al., 2010b) Amyotrophic Lateral Schlerosis (ALS) (Gil et al., 1993; Hanagasi et al., 2002; Pekkonen et al., 2004; Raggi et al., 2008)

Anaesthesis and sedation (Csepe et al., 1989; Heinke and Koelsch, 2005; Heinke et al., 2004; Koelsch et al., 2006; Astikainen et al., 2006; Yppärilä et al., 2002; Simpson et al., 2002; van Hooff et al., 1995, 1997) 424–458 (2012) 123 Neurophysiology Clinical / al. et Näätänen R. – Anhedonia (Giese-Davis et al., 1993) – Aphasia and stroke (Aaltonen et al., 1993; Alain et al., 1998; Auther et al., 2000; Becker and Reinvang, 2007; Csépe et al., 2001; Ilvonen et al., 2003, 2004; Jacobs and Schneider, 2003; Kohlmetz et al., 2001; Peach et al., 1992, 1993, 1994; Pettigrew et al., 2004a, 2005; Särkämö et al. 2010a,b; Wertz et al., 1998) – Asperger syndrome (Jansson-Verkasalo et al., 2003a, 2005; Korpilahti, 1995; Korpilahti et al., 2007; Kujala et al., 2005a, 2007, 2010; Lepistö et al., 2006, 2007) – Autism (Bomba and Pang, 2004; Ceponiene et al., 2003; Dunn et al., 2008; Ferri et al., 2003; Gomot et al., 2002; Kasai et al., 2005; Kemner et al., 1995; Kuhl et al., 2005; Lepistö et al., 2005, 2008; Näätänen and Kujala, 2011; Oram Cardy et al., 2005a,b; Roberts et al., 2008, 2011; Seri et al., 1999, 2007; Siegal and Blades, 2003) – Bipolar disorder (Hall et al., 2007, 2009; Takei et al., 2010) – Bipolar II disorder (Andersson et al., 2008) – Blindness (early) (Alho et al., 1993; Kujala et al., 1995a,b, 1997, 2000b) 999, 2000, 2001; Kohlmetz et al., 2001; Kotchoubey, 2007; – BrainKotchoubey lesions (Alain et et al. al., 2003, 1998; 2005 Alho; Mäkelä et al., 1994; et al., Auther 1998b; et Neumann al., 2000; andDeouell Kotchoubey, et al., 2000a,b; 2004; Hämäläinen Polo et al., 2002; et al., Reinvang 2007; Jacobs et al., and 2000; Schneider, Tarkka 2003; et al., Kaipio 2011; etWoods al., 1 and Knight, 1986); (see also Aphasia and Stroke) – Brainstem auditory processing syndrome (Kraus et al., 1993a) – CATCH-22 syndrome (Baker et al., 2005; Cheour et al., 1998a; Ceponiene et al., 2002) – Central auditory processing disorder (CAPD) (Bamiou et al., 2000) – Cerebellar degeneration (Moberget et al., 2008) – Childhood cancer and brain radiation (Järvelä et al., 2011; Lähteenmäki, 1999) – Chronic pain (Dick et al., 2003) – Chronic primary insomnia (Wang et al., 2001) – Cleft palate (different forms of) (Ceponiene et al., 2000, 2002; Cheour et al., 1998a, 1999) – Closed-head injury (Kaipio et al., 1999, 2000, 2001; Polo et al., 2002) – Cochlear-implant (CI) users (Groenen et al., 1996; Kelly et al., 2005; Kileny et al., 1997; Koelsch et al., 2004; Kraus et al., 1993b; Lonka et al., 2004; Nager et al., 2007; Ponton et al., 1996, 2000, 2009; Ponton and Don, 1995, 2003; Ponton and Eggermont, 2007; Roman et al., 2005; Salo et al., 2002; Sandmann et al., 2010; Wable et al., 2000) – Coma (Daltrozzo et al., 2007; Fischer et al., 1999, 2000, 2004, 2006a,b, 2008; Fischer and Luauté, 2005; Guérit et al., 1999; Kane et al., 1993, 1996, 2000; Kotchoubey et al., 2003; Laureys et al., 2005; Luauté et al., 2005; Morlet et al., 2000; Naccache et al., 2005; van der Stelt and van Boxtel, 2008; Vanhaudenhuyse et al., 2008) – Conduct disorder (Rothenberger et al., 2000) – Craniosynosthosis (Huotilainen et al., 2008) – Deafness to musical dissonance ( Acquired) (Brattico et al., 2003) – Dementia (Schroeder et al., 1995; Yokoyama et al., 1995; Osawa et al., 1996) – Depression (He et al., 2010; Iv et al., 2010; Kähkönen et al., 2007; Lepistö et al., 2004; Ogura et al., 1991, 1993; Takei et al., 2009) (continued on next page) 427 428 Table 1 (continued)

– Developmental dysphasia (Holopainen et al., 1997, 1998; Korpilahti and Lang, 1994) – Diabetes mellitus (late phase) (Vanhanen et al., 1996) – Distractibility (Escera et al., 2000a; Gumenyuk et al., 2004; Kaipio et al., 2000, 2001; Kilpeläinen et al., 1999; Mager et al., 2005) – Down’s syndrome (Diaz and Zurron, 1995; Lalo et al., 2005) – Drugs –Antipsychotics/Neuroleptics: – Clozapine (Horton et al., 2010; Schall et al., 1999; Umbricht et al., 1998) – Haloperidol (Kähkönen and Ahveninen, 2002; Kähkönen et al., 2001, 2002; Pekkonen et al., 2002) – Olanzapine (Korostenskaja et al., 2005) – Risperidone (Umbricht et al., 1999) – Alcohol: –(Hirvonen et al., 2000; Jääskeläinen et al., 1995, 1996a,b, 1998, 1999a; Kähkönen et al., 2001,2004,2005b; Kenemans et al., 2010) – Acetylcholine:

Tetrahydroaminoacridine (THA, Tacrine) (Engeland et al., 2002; Riekkinen et al., 1997) 424–458 (2012) 123 Neurophysiology Clinical / al. et Näätänen R. – (a) Muscarinic receptor antagonist: – Glycopyrrolate (Pekkonen et al., 2001a) – Scopolamine (Pekkonen et al., 2001a) – (b) Nicotinic receptor agonists: – Nicotine: (Baldeweg et al., 2006; Dulude et al., 2010; Engeland et al., 2002; Fisher et al., 2010; Harkrider and Hedrick, 2005; Inami et al., 2005, 2007; Knott et al., 2009; Martin et al., 2009; Pritchard et al., 2004) – AZD3480 (Dunbar et al., 2007) – Benzodiazepines: –(Kivisaari et al., 2007; Murakami et al., 2002) – Triatzolam (Nakagome et al., 1998) – Lorazepam (Rosburg et al., 2004) – GABA agonists/antagonists: – Bicuculline (Javitt et al., 1996) – Flumazenil (Smolnik et al., 1998) – Propofol (Heinke et al., 2004; Koelsch et al., 2006; Simpson et al., 2002) – Monoamines – Review: (Kähkönen and Ahveninen, 2002) – Adrenergic drugs: – ClonidineAtipamezole (Duncan (Mervaala and Kaye,et al., 1987;1993) Hansenne et al., 2003) – Noradrenergic agonist S 12024-4 (Missioner et al., 1999) – Histamine: – Antihistamines: Chlorpheniramine (Serra et al., 1996) – Dopamine: – Apomorphine (Hansenne et al., 2003) – Methamphetamine (Hosák et al., 2008) – Methylphenidate (Winsberg et al., 1993, 1997; Sawada et al., 2010; Verbaten et al., 1994) – Bromocriptine (Leung et al., 2007) – Domperidone (Leung et al., 2007) – Serotonin: – Serotonin (Kähkönen et al., 2005a; Oranje et al., 2008) – Acute tryptophan depletion (Kähkönen and Ahveninen, 2002) – Dimethyltryptamine (Heekeren et al., 2008) – Escitalopram (Oranje et al., 2008; Wienberg et al., 2010) – Psilocybin (Umbricht et al., 2003) – Neuropeptides/Hormones: – Vasopressin (Born et al., 1986, 1987a, 1998) – Oxytocin (Born et al., 1987a) – Cholecystokinin (Schreiber et al., 1995) – Desacetyl-alpha-MSH (Smolnik et al., 1999) – Hydrocortisone (Born et al., 1987a) – ACTH-related neuropeptides (Born et al., 1987b; Smolnik et al., 1999; Pietrowsky et al., 1990) – NMDA/Glutamate: – Ketamine (Ehrlichman et al., 2008; Heekeren et al., 2008; Javitt et al., 2000; Kreitschmann-Andermahr et al., 2001; Umbricht et al., 2002) – MK-801 (Javitt et al., 1996; Tikhonravov et al., 2010) – Phencyclidine (PCP) (Javitt et al., 1996) – Memantine (Korostenskaja et al., 2007) – Nitrous oxide (Pang and Fowler, 1999) – Glycine (Leung et al., 2008) – N-acetyl-cysteine (NAC; Lavoie et al., 2007) – Opiods: – Naltrexone (Jääskeläinen et al., 1998) – Opioid Dependence (Kivisaari et al., 2007; Näätänen, 2001) – Other/alternative therapies: – Hiba odour (Hiruma et al., 2002) – Purine nucleotides: – Adenosine (Hirvonen et al., 2000) – Caffeine (Rosburg et al., 2004) – Dyslexia (Ahissar, 2007; Ahissar et al., 2006; Alonso-Búa et al., 2006; Baldeweg et al., 1999; Bishop, 2007a,b; Blitz et al., 2007; Bonte et al., 2005, 2007; Bradlow et al., 1999; Bruder et al., 2011a,b; Corbera et al., 2006; Csépe,

2002; Czamara et al., 2011; Fisher et al., 2006; Froyen et al., 2008, 2009, 2010; Guttorm et al., 2005, 2001; Hämäläinen et al., 2008; Heim et al., 2000; Huttunen et al., 2007; Huttunen-Scott et al., 2008; Jansson-Verkasalo 424–458 (2012) 123 Neurophysiology Clinical / al. et Näätänen R. et al., 2010; Kujala, 2007; Kujala and Näätänen, 2001; Kujala et al., 2000a, 2001, 2003, 2006; Kurtzberg et al., 1995; Lachmann et al., 2005; Leppänen and Lyytinen, 1997; Leppänen et al., 1997, 1999, 2002, 2004, 2010; Lovio et al., 2010; Lyytinen et al., 2001, 2004, 2006, 2009; Maurer et al., 2003, 2009; Paul et al., 2006a,b; Pihko et al., 1999, 2008; Renvall and Hari, 2003; Richardson et al., 2003; Roeske et al., 2011; Schulte-Körne and Bruder, 2010; Schulte-Körne et al., 1998, 1999, 2001; Sebastian and Yasin, 2008; Shankarnarayan and Maruthy, 2007; Sharma et al., 2005, 2006, 2007, 2009; Stoodley et al., 2006; van Leeuwen et al., 2006) – Dysthymia (Giese-Davis et al., 1993) – Electromagnetic ﬁelds (Kwon et al., 2009, 2010) – Epilepsy (Boatman et al., 2008; Borghetti et al., 2007; Duman et al., 2008; Gene-Cos et al., 2005; Korostenskaja et al., 2010b; Liasis et al., 2000; Liasis et al., 2001; Liasis et al., 2006; Lin et al., 2007; Metz-Lutz and Philippini, 2006; Miyajima et al., 2011; Rosburg et al., 2005) – Frontal-lobe injury (Alho et al., 1994; Woods and Knight, 1986) – Herpes Simplex Encephalitis (Mäkelä et al., 1998a) – HIVHippocampus-Amygdala(Schroeder et al., 1994 Partial) Temporal-Lobe Resection (Hämäläinen et al., 2007) – Huntington’s disease (pathological enhancement of discrimination) (Beste et al., 2008) – Hyperkinetic disorder (Rothenberger, 1995) – Hypnosis (Jamieson et al., 2005; Kallio et al., 1999) – Insomnia (Wang et al., 2001) – Intellectual disability (Ikeda et al., 2004; Nakagawa et al., 2002; Kaga et al., 1999) – Introvert/extrovert personality (Sasaki et al., 2000) – Landau–Kleffner syndrome (Honbolygó et al., 2006; Metz-Lutz and Philippini, 2006) – Language-learning impairment (LLI) (Benasich and Tallal, 2002; Benasich et al., 2006; Bradlow et al., 1999; Choudhury and Benasich, 2011; Kraus et al., 1996; Kujala, 2007; Marler et al., 2002; Pihko et al., 2007; Shafer et al., 2005; Uwer et al., 2002; Weber et al., 2005) – Late talkers (Grossheinrich et al., 2010) – Learning-disabled children (Banai and Ahissar, 2005; Bradlow et al., 1999; Kraus et al., 1996) – Left sylvian infarct in neonate (Dehaene-Lambertz et al., 2004) – Locked-in patients (Ragazzoni et al., 2000) – Meditation (Srinivasan and Baijal, 2007) – Mental work load (Kramer et al., 1995) – Mental retardation (Holopainen et al., 1998; Ikeda et al., 2004; Kaga et al., 1999; Nakagawa et al., 2002; Yokoyama et al., 1995; Zurrón and Díaz, 1997) – Migraine (de Tommaso et al., 2004; Korostenskaja et al., 2010a; Valeriani et al., 2008) – Multiple schlerosis (MS) (Gil et al., 1993; Jung et al., 2006; Santos et al., 2006) – Narcolepsy (Naumann et al., 2001) ) – Neglect (Hemiﬁeld inattention) and extinction (Deouell et al., 2000a,b; Tarkka et al., 2011 – Noise and distraction (Bertoli et al., 2005; Gumenyuk et al., 2004; Mikkola et al., 2010; Shtyrov et al., 1998, 1999; Simoens et al., 2007) – Noise, different types of (Kozou et al., 2005) – Noise, occupational (Mäntysalo and Salmi, 1989) – Noise, occupational, long-term exposure (Brattico et al., 2005; Kujala and Brattico, 2009; Kujala et al., 2004) – Obsessive–compulsive disorder (Towey et al., 1994; Oades et al., 1996) – Obstructive sleep apnea syndrome (OSAS) (Gosselin et al., 2006) – Opioid dependence (Kivisaari et al., 2007) – Parkinson’s disease (Brønnick et al., 2010; Karayanidis et al., 1995; Pekkonen et al., 1995c; Vieregger et al., 1994) 429 – Perinatal asphyxia (Leipälä et al., 2011) (continued on next page) 430

Table 1 (continued)

– Perinatal intracerebral hemorrhage (Leipälä et al., 2011) – Persistent vegetative state (Boly et al., 2011; Fischer and Luauté, 2005; Kotchoubey, 2007; Kotchoubey et al., 2003, 2007, 2008; Laureys et al., 2005; Wijnen et al., 2007; Zarza-Lucianez et al., 2007) – Personality differences (Matsubayashi et al., 2008; Franken et al., 2005) – Phonological deﬁcit (Blitz et al., 2007) – Plagiocephaly (Huotilainen et al., 2008) .Nääe ta./Ciia erpyilg 2 21)424–458 (2012) 123 Neurophysiology Clinical / al. et Näätänen R. – Post-traumatic stress syndrome (PTSS) (Cornwell et al., 2007; Ge et al., 2011; Menning et al., 2008; Morgan and Grillon, 1999) – Psychosocial stress (Simoens et al., 2007) – Prematurely born (with very low birth weight, VLBW) infants (Bisiachi et al., 2009; Cheour-Luhtanen et al., 1996; Fellman and Huotilainen, 2006; Fellman et al., 2004; Gomot et al., 2007; Holst et al., 2005; Jansson-Verkasalo et al., 2004, 2010; Leipälä et al., 2011; Mento et al., 2010; Mikkola et al., 2007, 2010) – REM-sleep deprivation (Zerouali et al., 2010) – Schizophrenia (Ahveninen et al., 2006; Baldeweg et al., 2002, 2004; Banati and Hickie, 2009; Bodatsch et al., 2011; Braff and Light, 2004; Brockhaus-Dumke et al., 2005; Catts et al., 1995; Davalos et al., 2003, 2005; Devrim-Ücok et al., 2008; Ehrlichman et al., 2008, 2009; Fisher et al., 2008, 2011; Grzella et al., 2001; Hermens et al., 2010; Hirayasu et al., 1998, 2000, 2001; Inami et al., 2007; Jahshan et al., 2011; Javitt, 1993, 2000, 2009a,b; Javitt et al., 1996, 1998, 1999, 2000, 2008; Jessen et al., 2001; Kasai et al., 2001, 2002a,b,c, 2003a,b; Kathmann et al., 1995; Kaur et al., 2011; Kawakubo et al., 2006, 2007; Kiang et al., 2007, 2009; Kircher et al., 2004; Korostenskaja et al., 2005; Kreitschmann-Andermahr et al., 1999, 2001; Lavoie et al., 2007; Leitman et al., 2010; Light and Braff, 2005a,b; Luck et al., 2011; Magno et al., 2008; Matthews et al., 2007; Michie, 2001; Michie et al., 2002; Minami and Kirino, 2005; Morlet et al., 2000; Näätänen and Kähkönen, 2009; Niznikiewicz et al., 2009; Oades et al., 2006; Oknina et al., 2005; Park et al., 2002; Potts et al., 1998; Price et al., 2006; Rasser et al., 2011; Rissling et al., 2010; Salisbury et al., 2002, 2007; Sato et al., 1998, 2002; Schall et al., 1998, 1999, 2003; Schreiber et al., 1992; Shelley et al., 1991; Shin et al., 2009; Shinozaki et al., 2002; Stone et al., 2010; Thönnesen et al., 2008; Todd et al., 2008; Todd et al., in press; Todd et al., 2003, 2008; Turetsky et al., 2007, 2009; Umbricht et al., 1998, 1999, 2000, 2002, 2003, 2006; van der Stelt and Belger, 2007; Verbaten and van Engeland, 1995; Wible et al., 2001; Wynn et al., 2010; Yamasue et al., 2004; Youn et al., 2003; for a meta-analysis of 32 studies, see Umbricht and Krljes, 2005) – Serotonin transporter gene variants, individuals with (Sysoeva et al., 2009) – Sensorineural hearing loss (Oates et al., 2002; Stapells, 2002) – Sleep (Atienza et al., 1997, 2000, 2002; Atienza and Cantero, 2001; Campbell et al., 1991; Cheour et al. 2000,2002; Martynova et al., 2003; Nashida et al., 2000; Nittono et al., 2001; Ruby et al., 2008; Sallinen et al., 1994, 1996; Sculthorpe et al., 2009) – Sleep deprivation (Gumenyuk et al., 2010; Raz et al., 2001; Sallinen and Lyytinen, 1997; Salmi et al., 2005; Wang et al., 2001; Zerouali et al., 2010) – Socially withdrawn children (Bar-Haim et al., 2003) – Somatisation disorder (James et al., 1989) – Speciﬁc language impairment (SLI) (Barry et al., 2008; Bishop and McArthur, 2004; Friedrich et al., 2004; Korpilahti, 1995; Marler et al., 2002; Pihko et al., 2008; Rinker et al., 2007; Shafer et al., 2007; Uwer and von Suchodoletz, 2000; Uwer et al., 2002; Weber et al., 2005) – Stroke (see Aphasia) – Stuttering (Corbera et al., 2005; Wu et al., 1997) – Temporal-parietal brain lesions (Alain et al., 1998) – Thalamic infarction (Mäkelä et al., 1998b) – Tourette syndrome (TS) (Oades et al., 1996; Rothenberger et al., 2000; van Woerkom et al., 1988, 1994) – Tune deafness (Braun et al., 2008) – Velo-cardio-facial (DiGeorge) syndrome (Baker et al., 2005) R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 431

Table 2 THE MMN shows abnormally short auditory sensory-memory duration in:

– Alzheimer’s disease (Pekkonen et al., 1994) – Infants and children with cleft palate and CATCH-22 syndrome (Ceponiene et al., 2002; Cheour et al., 1997, 1998a, 1999) – Late talkers (Grossheinrich et al., 2010) – Chronic alcoholism (Grau et al., 2001; Marco-Pallares et al., 2007; Polo et al., 1999) – Normal ageing (Cooper et al., 2006; Grau et al., 2001; Jääskeläinen et al., 1999a; Pekkonen et al., 1996a) – Parents of children with speciﬁc language impairment (SLI) (Barry et al., 2008)

Table 3 language journals. Almost all these studies report, with only a few The MMN provides an index of increased backward masking (affecting speech exceptions, group-level results. perception) in:

– Chronic alcoholism (Ahveninen et al., 1999) 2. The MMN as an index of decreased auditory discrimination – Dyslexia (Kujala et al., 2003) – Speciﬁc language impairment (SLI) (Marler et al., 2002) accuracy (Table 1)

In several clinical conditions, the MMN amplitude is attenuated, duration, capacity and accuracy (Pekkonen et al., 1996a; Cooper usually indexing decreased behavioural discrimination accuracy et al., 2006; Polo et al., 1999; Grau et al., 2001). Furthermore, the (Javitt et al., 1998; Rabinowicz et al., 2000; Matthews et al., auditory MMN can even index auditory long-term memory traces 2007). This amplitude reduction was usually found by recording such as the language-specific memory traces, enabling one to cor- the MMN to simple frequency or duration changes in sinusoidal rectly perceive the speech sounds of the mother tongue and other tones, for instance, in schizophrenia (Todd et al., 2003; Javitt familiar languages (Dehaene-Lambertz, 1997; Näätänen, 2001; et al., 2000; Michie et al., 2000; see also Javitt et al., 1998). In Näätänen et al., 1997; Pulvermuller et al., 2004; Shtyrov and Pulver- schizophrenia, the duration MMN, in particular that to duration muller, 2007; Winkler et al., 1999c; Shtyrov and Pulvermüller, increment, was even more affected than that to frequency change 2007). This linguistic MMN subcomponent is usually left-hemi- (Michie, 2001; Michie et al., 2000; for a meta-analysis, see Umb- spherically predominant and generated posteriorly to the bilateral richt and Krljes, 2005). The frequency MMN is attenuated in ampli- MMN subcomponent for mere acoustic change (Näätänen et al., tude in several other clinical groups, too, such as children with 1997; Shestakova et al., 2002). developmental dysphasia (Holopainen et al., 1997, 1998; Kor- The extensive MMN literature of clinical studies can be classi- pilahti and Lang, 1994), dyslexia (Baldeweg et al., 1999; Renvall fied according to the kind of clinically useful information that and Hari, 2003; Kujala et al., 2003, 2006; Sebastian and Yasin, can be obtained by using the MMN. The MMN can index: 2008) and SLI (Mengler et al., 2005; Rinker et al., 2007). In children and adults with dyslexia, the MMN to frequency (1) Auditory discrimination accuracy (which is decreased in a change but not that to duration change was considerably attenu- number of clinical groups and affected by different drugs) ated in amplitude (Baldeweg et al., 1999; Kujala et al., 2006). Fur- (Table 1); thermore, in keeping with this profile of MMN-amplitude (2) shortened sensory-memory duration (and hence possibly reduction, the behavioural frequency but not duration discrimina- decreased general brain plasticity) (Table 2); tion was affected (Baldeweg et al., 1999). In addition, this (3) abnormal auditory perception; frequency-MMN deficit strongly correlated with the severity of (4) increased backward masking (Table 3); the reading problem. Subsequently, Lachmann et al. (2005) (5) abnormal involuntary attention switching, either too weak (Table 4) or too strong (Table 5); (6) cerebral grey-matter loss and other structural changes (Table 6); (7) pathological brain excitation/excitability state (Table 7); MMN (8) cognitive and functional decline (Table 8); MMN (9) the level of consciousness (Table 9); (10) the progression of illness (Table 10); (11) future clinical condition (prognosis) (Table 11); Amplitude (12) genetic disposition to certain disorders (Table 12); and (13) recovery/improvement as a function of time or treatment Time (Table 13). Moreover, recent studies using the vMMN (Iijima et al., 1996; Lorenzo-Lopez et al., 2004; Tales and Butler, 2006; Left Right Tales et al., 2002a,b; Kenemans et al., 2010; Tales and Butler, hemisphere hemisphere 2006; Tales et al., 2002a, 2008; Maekawa et al., 2011; Froyen et al., 2010; Chang et al., 2010; Qiu et al., 2011; Horimoto et al., 2002; Hosák et al., 2008; Kremlácˇek et al., 2008, Ver- 3-5 days baten et al., 1994; Tales and Butler, 2006; Tales et al., 10 days 2008; Fisher et al., 2010; Urban et al., 2008; for a recent 3 months review, see Kimura et al., 2011) and sMMN (Restuccia et al., 2007; Akatsuka et al., 2005, 2007; Näätänen, 2009) Fig. 3. MMN shows how frequency discrimination recovered to normal levels have also provided clinically important results. within 3 months from a stroke onset (averaged data of 8 patients). This MMN recovery was slower for right-ear stimulation (with the main part of the auditory input going to the damaged left hemisphere) (curves on the left) than with left-ear The present review aims at covering all clinical MMN studies in stimulation (curves on the right). During the MMN recording, the patient was the auditory modality published in refereed international English- watching video-films. Adapted from Ilvonen et al. (2003). 432 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458

Table 4 The MMN shows decreased involuntary attention switching in:

– Alcohol intoxication (Jääskeläinen et al., 1995, 1996a,b, 1999b) – Schizophrenia (Baldeweg et al., 2002; Catts et al., 1995; Matthews et al., 2007; Oades et al., 2006; Sato et al., 2003) – Closed-head injury (Polo et al., 2002) – Frontal-lesion patients (Alho et al., 1994) – Right-hemisphere damage (unilateral neglect) (Deouell et al., 2000a,b) – Children with major depression (Lepistö et al., 2004)

showed that the MMN-attenuation profile involving frequency and see also Näätänen and Kujala, 2011) that in autistic children of consonant–vowel (CV) syllable changes could disentangle different 7–9 years, the MMNm peak latency for tone-frequency and vowel diagnostic subgroups of developmental dyslexia from each other. changes was delayed, in comparison with that of normally devel- Moreover, Shafer et al. (2005) and Uwer et al. (2002) found that oping children of the same age, and, further, that this delay was SLI children had attenuated MMN amplitudes for syllable change. considerably increased if the child also suffered from a delay in lin- Using the new multi-feature MMN ‘optimum’ paradigm (Näätä- guistic development. This might contribute to their speech-under- nen et al., 2004), permitting one to obtain five different MMNs in standing difficulties and hence to their social isolation. In addition, 15 min, Lovio et al. (2010) found a widespread auditory discrimi- in the Asperger syndrome, a condition belonging to the autism nation deficit in children at risk for dyslexia. Their MMNs for con- spectrum, the duration-increment MMN of children of about sonant, vowel, vowel duration and intensity changes of syllables 9 years of age was attenuated in amplitude over the left hemi- were diminished in amplitude. By contrast, Kujala et al. (2006), sphere, whereas it was enhanced over the right hemisphere also using the multi-feature paradigm, found that in adult dyslex- (Lepistö et al., 2006). Moreover, in this syndrome, the MMN ampli- ics, the frequency MMN was attenuated, whereas their location tude was also attenuated for changes in prosody (Kujala et al., MMN was in fact enhanced in amplitude. This shows that not all 2005a). Recently, a shortened MMN peak latency for frequency changes in central auditory processing in dyslexia signify deterio- change and increased MMN amplitudes for duration and gap rated discrimination. However, the MMN data may also suggest changes were found in Asperger adults by Kujala et al. (2007). the presence of cognitive deficiencies associated with dyslexia, at In addition, the MMNs for frequency and duration changes were least deficient phonotactic processing (Bonte et al., 2007). attenuated in amplitude in patients with a left-hemispheric stroke Furthermore, in children with autism, the MMN to duration (Csépe et al., 2001; Aaltonen et al., 1993; Auther et al., 2000; Ilvo- change (Lepistö et al., 2005) and the MMNm to speech-sound nen et al., 2001, 2003; Pettigrew et al., 2005). These two MMNs change (Kasai et al., 2005; for recent corroborating results, see Rob- showed somewhat different time courses of post-stroke recovery, erts et al., 2011) were attenuated in amplitude. In addition, the however (Ilvonen et al., 2003)(Fig. 3). Importantly, the MMN MMN to CV-syllable change was prolonged in peak latency but recovery indexed that of speech perception, there being a correla- only in those children who preferred non-speech analogy signals tion between the duration-MMN amplitude and the Boston Diag- to speech (Kuhl et al., 2005). By contrast, the MMN to frequency nostic Aphasia Examination Speech-Comprehension Test from change was enhanced in amplitude (Lepistö et al., 2005; Ferri 10 days to 3 months post-stroke onset (Ilvonen et al., 2003). Subse- et al., 2003) and shortened in peak latency (Gomot et al., 2002) quently, Särkämö et al. (2010b) found that the MMNm recovery in autistic children, consistent with their better-than-average pitch and behavioural discrimination could be expedited with music discrimination and musical abilities (Bonnel et al., 2003; Heaton and speech stimulation. In addition, the MMNm recovery corre- et al., 2001). Very recently, it was found by Roberts et al. (2011;

Fig. 4. Scatterplot of MMN amplitude at Fz versus age, for schizophrenia patients (open circles; dashed regression line) and normal control subjects (solid circles; solid regression line) (From Kiang et al., 2009). R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 433

Table 5 F3 Fz F4 The MMN shows increased involuntary attention switching in:

– Closed head injury (Kaipio et al., 2000) 0.5 s – Ageing (Gaeta et al., 2001) – Alcoholism (Ahveninen et al., 2000b; Polo et al., 2003) – Children with major depression (Lepistö et al., 2004) – Sleep disorder (Gumenyuk et al., 2010) MMN

4.5 s lated with the recovery of behavioural speech-sound discrimination. Younger Males (mean 22 yrs) In epilepsy, the MMN is also affected. The MMN peak latencies Older Males (mean 59 yrs) were abnormally long or no clear MMN response could be found for speech (but could for tones) in patients with benign childhood Fig. 5. MMN to frequency change (700 ? 600 Hz) was normal in older males with epilepsy with contra-temporal spikes (Boatman et al., 2008; Du- an inter-stimulus interval (ISI) of 0.5 s but was considerably attenuated, relative to man et al., 2008). Patients with such spikes with atypical features that of younger males, with an ISI of 4.5 s, indexing faster auditory sensory-memory trace decay with aging (From Pekkonen et al., 2001a). and learning difficulties also exhibit attenuated MMN amplitudes (Metz-Lutz and Philippini, 2006). In addition, in the Landau–Kleff- ner syndrome, an MMN was obtained for phoneme change but not level of performance in different cognitive tasks is gradually de- for prosodic change (Honbolygó et al., 2006). By contrast, Miyajima creased (Cansino, 2009). There is also evidence linking these et al. (2011), studying adult patients with temporal-lobe epilepsy, MMN changes to cognitive deterioration (Kisley et al., 2005). Even found that at fronto-central sites, their MMN amplitude for tone- in healthy elderly persons, the MMN to a short gap in the middle of frequency change was enhanced, interpreted by the authors as a tone (Desjardin et al., 1999) was attenuated in amplitude, index- reflecting frontal-lobe hyperexcitability to compensate for tempo- ing their increased difficulties in correctly perceiving rapid speech- ral-lobe dysfunction indexed, according to the authors, by pro- sound patterns such as consonants (Bertoli et al., 2002, 2005). longed MMN peak latencies in patients relative to controls at The MMN and MMNm are also affected by different drugs both fronto-central and mastoid sites, consistent with previous re- (Table 1). ports (Lin et al., 2007; Borghetti et al., 2007). Furthermore, in the Miyajima et al. (2011) temporal-lobe epilepsy patients, the fron- to-centrally recorded MMN persisted longer than that in controls, 3. The MMN as an index of shortened sensory-memory duration consistent with the results of Gene-Cos et al. (2005) who suggested (Table 2) that a prolonged MMN duration might point to difficulty in ‘the closure mechanism of the MMN process’. The MMN elicitation depends on the presence of the sensory- The MMN can also be used to evaluate the possible central- memory trace representing the preceding stimuli and their regu- auditory-processing damage associated with a very premature larities at the moment of the delivery of a deviant stimulus (for a birth. Studying very prematurely born children (mean gestational review, see Näätänen et al., 2007). Hence, by gradually prolonging age 29 weeks) with a very low birth weight (mean 1115 g), Jans- the inter-stimulus interval (ISI), the MMN eventually vanishes, son-Verkasalo et al. (2003b, 2004) found that their MMN ampli- which enables one to assess sensory-memory duration in audition tude for consonant change at 4 years (from conception) was (a potential general index of brain plasticity; Näätänen and Kreegi- smaller than that of controls and, further, that the absence of the puu, 2011). MMN at 4–5 years of age predicted naming difficulties at 6 years. In young healthy adults, the trace duration, as estimated in this However, if the MMN amplitude nevertheless was normal at manner, approximates 10 s (Böttcher-Gandor and Ullsperger, 4 years, then there were no naming difficulties at 6 years. 1992; Sams et al., 1993), but is gradually shortened with ageing. Moreover, the MMN enables one to also determine central- With short ISIs such as 0.5 s, the MMN amplitude for frequency auditory-processing damage caused by long-term exposure to loud change was very similar in elderly (mean 59 years) and young occupational noise. In workers exposed to such a noise for 2– (mean 25 years) male subjects, suggesting that the memory-trace 17 years, the MMN to syllable change was attenuated in amplitude formation for sounds and, thus, perception (see Näätänen and Win- (even though their peripheral hearing was not affected), in keeping kler, 1999) were not affected by ageing. By contrast, with a stimu- with their increased speech-perception problems (Kujala et al., lus-onset asynchrony (SOA) of 4.5 s, the MMN amplitude of the 2004). In addition, the Brattico et al. (2005) MMN data showed that elderly was much more attenuated than that of the young (Pekko- long-term exposure to occupational noise altered the cortical orga- nen et al., 1996a)(Fig. 5), indicating that in ageing, the auditory nisation of speech-sound processing in these workers (in the ab- sensory-memory duration is shortened. Moreover, corresponding sence of peripheral hearing loss) but did not alter that of non- MMN data for frequency change in chronic alcoholics suggest speech processing. In addition, the discrimination of auditory stim- accelerated age-related shortening of the memory-trace duration uli was generally slowed down, as indexed by prolonged MMN- in these patients (Polo et al., 1999; see also Grau et al., 2001). peak latencies. The memory-trace duration is particularly short in patients Furthermore, by using the MMN, one can also monitor age-re- with Alzheimer’s disease, of course. Their MMN was, however, nor- lated changes in central auditory processing (Fabiani et al., 2006; mal with short ISIs (0.5 s), suggesting normal memory-trace for- Woods, 1992; Woods and Clayworth, 1986; Jääskeläinen et al., mation/perception (see Näätänen and Winkler, 1999) even in 1999a; Kiang et al., 2009; Cooper et al., 2006; Gaeta et al., 1998, these patients, but absent at long ISIs (3 s) (Pekkonen et al., 2001, 2002; Kisley et al., 2005; Pekkonen, 2000; Pekkonen et al., 1994), indexing their extremely short sensory-memory duration. 1995a, 1996a). A particularly convincing characterisation of the Interestingly, this data pattern is orthogonal to that observed in pa- MMN–age relationship was provided by Kiang et al. (2009) who tients with schizophrenia in whom no normal-amplitude MMN can carefully plotted the MMN and P3a amplitudes for tone-duration be obtained in any condition, not even with very short ISIs and increment across adulthood for 147 normal subjects and 253 wide frequency deviations (Javitt et al., 1998). This suggests that schizophrenic patients (Fig. 4). Along with an increasing age, the their memory-trace formation, and hence auditory perception 434 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458

Table 6 4. The MMN as an index of abnormal auditory perception The MMN correlates with structural changes in:

– Schizophrenia (Hirayasu et al., 2000, 2001; Kasai et al., 2003a,b; Park et al., In the foregoing, it was already mentioned that in patients with 2002; Rasser et al., 2011; Salisbury et al., 2002, 2007; Yamasue et al., 2004; schizophrenia, no normal-size MMN can be obtained irrespective Youn et al., 2003; for a review, see van der Stelt and Belger, 2007) of experimental manipulations, not even with very short SOAs (Ja- – Cleft palate (children with) (Ceponiene et al., 1999) vitt et al., 1998). This suggests a fundamental abnormality in memory-trace formation, and thus in auditory perception (see Näätänen and Winkler, 1999), in these patients (Näätänen and Kähkönen, 2009). Moreover, a number of further studies in patients with schizophrenia (Oades et al., 1996; Hirayasu et al., 1998; Schall et al., 1999; Youn et al., 2003; Kasai et al., 2002a,b) showed that the frequency-MMN amplitude deﬁcit tends to correlate with the severity of auditory hallucinations. In addition, Youn et al. (2003) found that the abnormality of the left–right MMN-amplitude -4 µV asymmetry for frequency change of a tone correlated with the positive scores of the positive-and-negative syndrome scale (PANSS), in particular with its hallucination subscale. For corroborating MMNm data, see Thönnesen et al. (2008). Moreover, Fisher et al. F4 (2008) found a smaller duration-MMN amplitude in patients with +1 µV 350 ms auditory hallucinations than in those with no hallucinations. In addition, the MMN amplitude for intensity change was also smaller 350-ms ISI in hallucinating than non-hallucinating patients. Furthermore, 700-ms ISI both patient groups showed an attenuated MMN amplitude for fre- 1400-ms ISI quency change. This MMN deﬁciency however was differently distributed in frontal areas in the two patient groups. Fig. 6. MMN for tone-frequency change was of normal amplitude in 7–9 year old children with oral clefts when a short ISI (350 ms) was used but was considerably smaller than that of controls when the ISI was prolonged to 700 ms and to 1400 ms, 5. The MMN as an index of increased backward masking suggesting an accelerated sensory-memory trace in patients (From Ceponiene et al., 1999). (Table 3)

Auditory masking refers to any observation such that informa- (Näätänen and Winkler, 1999), is abnormal, consistent with clini- tion in a test auditory stimulus is reduced by the presentation of cal observations (for a review, see Näätänen and Kähkönen, another (masking) auditory stimulus (Massaro, 1973). The percep- 2009). By contrast, their sensory-memory duration, as judged from tion of sequentially presented auditory stimulation may be hin- the ISI–MMN relationship in these patients, is unaffected (Javitt dered by backward masking, with a sound preventing the et al., 1998; see also March et al., 1999). perception of an immediately preceding sound, apparently by The auditory memory-trace duration shows developmental affecting its memory-trace formation (Massaro, 1975), in particular changes also in the beginning of the life, but to the opposite direc- when the later sound is stronger in intensity than the preceding tion, of course, being only 0.5 s in newborns, as judged from MMNs sound (see Hawkins and Presson, 1986). Massaro (1975) concludes recorded during sleep (Cheour et al., 2002b), but is thereafter grad- that backward masking occurs because the masking stimulus ually prolonged during infancy and childhood. By using the Grau essentially overwrites the auditory image of the test stimulus et al. (1998) MMN paradigm for effectively determining the sen- and, therefore, terminates the perceptual processing of the test sory-memory duration, Glass et al., 2008a,b) found that this dura- stimulus. tion was about 1–2 s at the age of 2–3 years but was prolonged to Backward masking can be assessed both behaviourally and by 3–5 s by the age of 6 years (see also Gomes et al., 1999). using the MMN: a masker presented shortly after each stimulus This memory-trace duration is affected in some child patient of the oddball paradigm causes the MMN to deviants and the subgroups, too. In children of 7–8 years with cleft palate, the MMN ject’s ability to discriminate them behaviourally to disappear in for tone-frequency change was of normal amplitude with short parallel (Winkler and Näätänen, 1994; Winkler et al., 1993). By SOAs (350 and 700 ms), whereas the prolongation of the SOA to using this type of paradigm, it was found by Kujala et al. (2003) 1400 ms attenuated this amplitude much more than it attenuated that backward masking is enhanced in dyslexic individuals. In- that of control children (Ceponiene et al., 1999)(Fig. 6). Further- creased backward masking, as reflected by MMN-data pattern, more, an analogous data pattern was obtained in small infants with was also reported in chronic alcoholism (Ahveninen et al., 1999). cleft palate, which was even more abnormal in children with the Importantly, this MMN deficit in chronic alcoholism correlated Catch-22 syndrome (Cheour et al., 1997, 1998a, 1999). with the amount of their alcohol consumption and predicted their In addition, a short auditory sensory-memory duration even in working-memory performance deficit (Ahveninen et al., 1999). normal healthy children is associated with delayed linguistic development. Late talkers had a shorter sensory-memory duration 6. The MMN as an index of abnormal involuntary attention than that of controls, as judged from MMN data for tone-frequency switching: either too weak (Table 4) or too strong (Table 5) change as a function of the ISI recorded at the age of 4 years 7 months, even though the language development was normal by As already mentioned, in addition to its auditory-cortex gener- that age (Grossheinrich et al., 2010). ators, the MMN also has a frontal (usually right-hemispherically Furthermore, the Barry et al. (2008) recent MMN data for tone- predominant (Giard et al., 1990)) generator contributing, together frequency change as a function of the ISI suggested a shorter audi- with the auditory-cortex MMN generator (see Fig. 2), to the fron- tory sensory-memory duration in parents of children with SLI than tally and centrally recorded MMN amplitudes (Deouell, 2007; that in parents of normally developing children. Giard et al., 1990; Rinne et al., 2000). The frontally recorded MMN amplitude is considerably attenuated, whereas the R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 435

Table 7 A similar (but transient) dampening of involuntary attention The MMN indexes pathological brain excitation/excitability in: switching can also be observed in acute alcohol intoxication – Alcoholism (Ahveninen et al., 2000a; Pekkonen et al., 1998) (Jääskeläinen et al., 1995, 1996a,b, 1998, 1999b). Even a moderate – ALS (with bulbar signs) (Pekkonen et al., 2004) dose of ethanol selectively weakened the frontal-MMN process, – Chronic primary insomniacs (Wang et al., 2001) whereas the supratemporal MMN process (with most of it being – Epilepsy (Miyajima et al., 2011) separately observable in nose-referenced mastoid recordings) – Huntington’s disease, Syndromatic Phase (Beste et al., 2008) – Major depression, children with (Lepistö et al., 2004) was unaffected (Jääskeläinen et al., 1996b)(Fig. 2). Furthermore, – Pediatric migraine (Valeriani et al., 2008; see, however, de Tommaso et al., ethanol abolished the performance decrement caused by auditory 2004) distraction in a visual task (Jääskeläinen et al., 1999b), i.e., damp- – Schizophrenia (pathological microglia activation, indexing local disease ened the effects of auditory distracting stimuli on brain mecha- process and neuronal death) (Banati and Hickie, 2009) – Stuttering (Corbera et al., 2005; see also Wu et al., 1997) nisms controlling for environment-driven involuntary attention switches in audition (Escera et al., 2000a; Escera and Corral, 2007; Jääskeläinen et al., 1999b). This explains, in part, why alcohol increases accident risk in traffic (cf. Näätänen and Summala, Table 8 1976). In addition, the Alho et al. (1994) MMN data suggested de- The MMN indexes cognitive and functional decline in: creased involuntary attention switching to auditory changes as a consequence of a frontal-lobe lesion (see Table 4). – Ageing (Gaeta et al., 2002; Kisley et al., 2005; Schroeder et al., 1995) – Alzheimer’s disease and dementia (Engeland et al., 2002; Schroeder et al., Conversely, the frontal involuntary attention-switching mecha- 1995) nism may also become too sensitive, in this way disturbing pri- – Amyotropic lateral schlerosis (ALS) (Raggi et al., 2008) mary-task performance (Table 5). In patients with closed head – Autism (Seri et al., 2007; Roberts et al., 2008, 2011; Näätänen and Kujala, injury suffering from dramatically increased distractibility, this 2011) – CATCH-22 syndrome: children (for a review, see Ceponiene et al., 2002), mechanism is abnormally sensitive, judging from their very large adults (Baker et al., 2005) frontally recorded MMN amplitudes at the presence of normal – Children with cleft palate (Ceponiene et al., 2002) mastoid-recorded (nose-referenced) MMN amplitudes (Kaipio – Chronic alcoholism (Polo et al., 1999) et al., 1999, 2000) (cf. Fig. 2). A subsequent study (Polo et al., – Coma, survivors of (Kane et al., 2000) 2002) did not confirm this finding, however, which might result – Developmental dysphasia (Holopainen et al., 1997, 1998; Korpilahti and Lang, 1994) from the abnormally fast vigilance decrement of these patients – Diabetes mellitus (advanced phase) (Vanhanen et al., 1996) (Kaipio et al., 2001). – Down’s syndrome (Lalo et al., 2005) Moreover, in children suffering from major depression, short- – Dyslexia (Bonte et al., 2007) ened MMN latencies and increased P3a amplitudes (an index of – Epilepsy (Boatman et al., 2008; Liasis et al., 2000, 2001, 2006; Metz-Lutz and Philippini, 2006) the occurrence of an attention switch; Squires et al., 1975; Escera – HIV (Schroeder et al., 1994) et al., 2000a) might be associated with these patients’ increased – Intellectual disability (Ikeda et al., 2004; Nakagawa et al., 2002) distractibility and difficulties in concentrating on task perfor- – Multiple schlerosis (MS) (Gil et al., 1993; Jung et al., 2006; Santos et al., 2006) mance, as shown by the Lepistö et al. (2004) MMN data for CV-syl- – Parkinson’s disease (Brønnick et al., 2010; Karayanidis et al., 1995; Pekkonen lable change. Analogous MMN evidence for increased et al., 1995c) – Persistent-vegetative-state (PVS), survivors of (Kotchoubey, 2007; Wijnen distractibility was also obtained in chronic alcoholism (Ahveninen et al., 2007; van der Stelt and van Boxtel, 2008; Zarza-Lucianez et al., 2007) et al., 2000a,b) and ageing (Gaeta et al., 2001). – Preterm children with very low birth weight (Jansson-Verkasalo et al., 2004, 2010) – REM-sleep deprivation (Zerouali et al., 2010) – Schizophrenia (e.g., Baldeweg et al., 2004; Hermens et al., 2010; Kawakubo 7. The MMN attenuation caused by cerebral grey-matter loss or et al., 2007; Light and Braff, 2005a,b; Rasser et al., 2011; Toyomaki et al., other structural change (Table 6) 2008) – Sleep deprivation (Raz et al., 2001) The MMN is also attenuated by structural damage in the central – Stroke and aphasia (Auther et al., 2000; Ilvonen et al., 2003, 2004; Pettigrew auditory system and even elsewhere in the brain. In schizophrenia et al., 2005; Särkämö et al., 2009, 2010a,b; Wertz et al., 1998) – Velo-cardio-facial syndrome (Baker et al., 2005) patients, a gradual loss of the left-hemisphere temporal grey-matter volume has been observed, which was reflected by attenuated MMN amplitudes for frequency change (Hirayasu et al., 1998). mastoid-recorded MMN (with nose reference) is unaffected in pa- More recently, Salisbury et al. (2007) found, by using magnetic res- tients with schizophrenia (Baldeweg et al., 2002, 2004; Sato et al., onance imaging (MRI), that at first hospitalisation, schizophrenia 1998, 2002), suggesting dampened involuntary attention switch- patients’ left Heschl-gyrus volume did not differ from that of bipo- ing to auditory change and hence probably accounting for lar-disorder patients and normal controls and, further, that this some of their withdrawal (negative) symptoms, as proposed by was reflected by similar MMN amplitudes for tone-frequency Näätänen and Kähkönen (2009) (cf. Fig. 2). Consistent with this, change. However, in an 18-month follow-up, this volume was re- Matthews et al. (2007) found that the MMN at fronto-central elec- duced in schizophrenia patients only, and the magnitude of the trodes for change in the interaural time difference, one of the two Heschl’s gyrus volume reduction in these patients strongly corre- main cues for sound lateralisation, was attenuated in amplitude in lated with the parallel attenuation of their MMN amplitude. Also, patients with schizophrenia. This result was attributed by the Yamasue et al. (2004) found that in schizophrenia patients, MMNm authors to patients’ impaired ability to use temporal cues to sound deficiency for phoneme change in the left hemisphere correlated lateralisation in natural settings, a deficit probably affecting, with the magnitude of the left planum-temporale grey-matter loss, according to the authors, their ability to control the focus of atten- suggesting that these structural abnormalities may underlie the tion effectively. The dampened bottom-up attention-switching functional abnormalities of fundamental language-related process- function in schizophrenia patients might also contribute to their ing in these patients. In addition, very recently, Rasser et al. (2011) gradual cognitive decline because of the resulting reduction of cog- found that MMN-amplitude attenuation for frequency change in nitive and social stimulation (cf. sensory deprivation) (Näätänen patients with schizophrenia (reflecting their impaired day-to-day and Kähkönen, 2009). For further supportive MMN evidence, see functioning level) correlated with the magnitude of grey-matter Hermens et al. (2010). loss in the left Heschl’s gyrus as well as in the motor and executive 436 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458

Fig. 7. Post Hoc correlation analyses of reduced gray matter measures – averaged over specific gyri defined on the Deformed ‘‘Montreal Neurological Institute’’ brain template – with reduced frequency mismatch negativity (MMN) peak amplitudes recorded at fronto-central electrodes in control subjects (blue) and schizophrenia patients (Red). Solid regression lines indicate significant correlation (P < .05, uncorrected) (From Rasser et al., 2011). R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 437

Table 9 Consistent with the Pekkonen et al. (2004) above-reviewed ALS The MMN can be used to on-line monitor the patient’s level-of-consciousness in: results, at late, symptomatic, stages of Huntington’s disease, the – Anaesthesis: depth and recovery (Heinke et al., 2004; Heinke and Koelsch, MMN to frequency change was considerably enhanced in ampli- 2005; Koelsch et al., 2006; Yppärilä et al., 2002) tude, along with behavioural improvement in an auditory signal- – Coma (Kane et al., 1993, 1996, 2000; Fischer and Luauté, 2005; Luauté et al., detection task, compared to the control group and Huntington 2005; Morlet et al., 2000; Naccache et al., 2005) patients in the early, presymptomatic, stage (Beste et al., 2008). – Persistent vegetative state (Kotchoubey, 2007; Kotchoubey, 2007; Wijnen et al., 2007) This paradoxical effect, as contrasted to effects of neurodegenerative diseases in general, was attributed by the authors to toxic-level overexcitability of the NMDAR system (due to increased glutamate concentrations). regions of the frontal cortex while MMN reduction for duration In addition, very importantly, Banati and Hickie (2009); (see deviants also correlated with grey-matter loss in the right Heschl’s also Banati, 2003) found in patients in schizophrenic psychosis that gyrus (Fig. 7). MMN deficiency for frequency and duration deviants indexed It is possible that the MMN deficiency observed in schizophre- pathological microglia (the main constituent of the brain’s innate nia can be linked to grey-matter loss occurring in these patients. immune system) activation, reflecting local ‘disease activity’ and Olney and Farber (1995) suggested that N-methyl-D-aspartate neuronal death in these patients. The authors emphasised that this (NMDA)-receptor (NMDAR) hypofunction accounts for structural microglia activation is not a diagnostically specific pathological brain changes in these patients. The data reviewed by the authors process but rather that this activation can serve as a generic mar- showed, among other things, that a repeated treatment with an ker that relates more directly to disease progression and may be NMDA antagonist during a 3- to 4-day period induces a pattern correlated with other meaningful illness measures such as the of neurodegenerative changes distributed over several corticolim- MMN. bic brain regions. The authors concluded that these and other find- Pathological brain excitation, as suggested by increased MMN ings signify that the persistent suppression of the NMDAR function amplitudes, can also be found in temporal-lobe epilepsy (Miyajima lasting for a few days only can lead to relatively subtle but perma- et al., 2011; Gene-Cos et al., 2005) even though MMN peak laten- nent structural changes that are distributed over the neocortical cies may be prolonged, a possible sign of central-auditory-process- and limbic brain regions in a pattern roughly similar to the pattern ing disturbance in these patients (Miyajima et al., 2011). of structural changes observed in schizophrenia (Bogerts, 1993). Pathological excitation might also explain some results in pa- The fact that the MMN is an index of NMDAR dysfunction would tients suffering from post-traumatic stress disorder (PTSD), who then explain the relationship between MMN deficiency and grey- showed considerably elevated MMN amplitudes for frequency matter loss (Salisbury et al., 2007; Rasser et al., 2011). change (Morgan and Grillon, 1999; Ge et al., 2011), whereas Men- The MMN can also reflect structural changes (craniofacial mal- ning et al. (2008) could not confirm these results. formations) associated with cleft palate (for a review, see Ceponi- ene et al., 2002). In children with cleft palate of 7–9 years of age, it was found by Ceponiene et al. (1999) that the more posteriorly 9. The MMN as an index of cognitive and functional delimited the cleft was, the smaller was the MMN amplitude for deterioration (Table 8) tone-frequency change and the more severe was the cognitive defect. Baldeweg et al. (2004) found a relationship between the frontally recorded duration-increment MMN deficit and impairments 8. The MMN as an index of pathological brain excitation/ in memory functions of patients with schizophrenia, which are evi- excitability (Table 7) dent in these patients (Sullivan et al., 1995; Jahshan et al., 2010). Subsequently, Light and Braff (2005a) obtained a strong correlation In some conditions, an increased MMN amplitude appears to in- between the global-assessment-of-functioning (GAF) ratings and dex pathologically increased central nervous system (CNS) or cen- the fronto-central MMN amplitude for tone-duration prolongation tral-auditory-system excitation/excitability. For example, in in these patients. Furthermore, the MMN amplitude was highly abstinent chronic alcoholics, the MMN for frequency change was predictive of the patients’ level of independence in their domestic abnormally enhanced in amplitude, which might be associated life. This correlation was replicated in longitudinal studies 1– with their increased distractibility (Ahveninen et al., 2000a,b). In 2 years later, indicating a stable relationship (Light and Braff, addition, in patients with persistent developmental stuttering, 2005b). Subsequently, the relationship between the duration- the MMN to phonetic contrasts was strongly enhanced in ampli- increment MMN amplitude and patients’ functional status was also tude, whereas that to simple tone contrasts (both frequency and found by Kawakubo et al., 2007; social skills acquisition) and by duration) was unaffected (Corbera et al., 2005). This might index Toyomaki et al. (2008; executive functioning). (For corroborating abnormal excitability located specifically in the speech-sound results with the frequency-MMN amplitude, see Rasser et al. MMN generation region (Näätänen et al., 1997). This MMN finding (2011).) Interestingly, this relationship for duration-increment correlated with the patients’ subjective appraisal of their speech- MMN amplitude is also present in normal healthy subjects (Light production problems. These results might indicate that proper et al., 2007). For previous analogous results with the frequency- speech production requires the presence of stable sensory refer- MMN amplitude in normal healthy subjects, see Bazana and Stel- ence provided by the speech-sound-specific memory traces of mack (2002) and Beauchamp and Stelmack (2006), and with the the left temporal cortex (Näätänen, 2001; Näätänen et al., 1997; MMN amplitude for syllable change in children, Liu et al. (2007). Shestakova et al., 2003). For recent corroborating results, see Troche et al. (2009, 2010). In the same vein, an MMNm enhancement to tone-duration In a similar vein, Jung et al. (2006) found that the MMN-ampli- decrement found by Pekkonen et al. (2004) in patients with ALS tude attenuation for duration decrement in patients with MS in- with bulbar signs might result from cortical overactivity of excit- dexed cognitive decline occurring in part of these patients, being atory neurotransmitter glutamate observed in ALS (Rowland and larger in magnitude for greater cognitive loss. For corroborating re- Shneider, 2001). Raggi et al. (2008), however, found MMN-ampli- sults, see Gil et al., 1993) and Santos et al. (2006). Furthermore, tude decrement for frequency change in non-demented patients MMN (MMNm) deficit indexes cognitive decline also in chronic with sporadic ALS. alcoholism (Polo et al., 1999, 2003), stroke (Ilvonen et al., 2003; 438 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458

Table 10 tude increase when the patient crossed the boundary from uncon- The MMN can be used to monitor the patient’s state in: sciousness to consciousness. In addition, in locked-in patients, the – Stroke: recovery (Ilvonen et al., 2003; Särkämö et al., 2010a,) frequency MMN can give evidence for the presence of at least some – Schizophrenia: illness progression (Banati and Hickie, 2009; Bodatsch et al., level of consciousness (Ragazzoni et al., 2000). Moreover, the MMN 2011; Javitt et al., 2000; Oades et al., 2006; Oknina et al., 2005; Salisbury can also index anaesthesia depth (for a review, see Heinke and et al., 2007; Umbricht and Krljes, 2005; Umbricht et al., 2003, 2006) and Koelsch, 2005): a small-amplitude MMN was recorded to fre- temporary recovery (Shinozaki et al., 2002) quency change during deep sedation, whereas in complete unconsciousness, this MMN was absent. It, however, returned while the patient was recovering from anaesthesia (Heinke et al., 2004). Interestingly, an MMN type of response was also elicited by mu- Särkämö et al., 2009, 2010a,; Auther et al., 1998, 2000; Pettigrew sic-syntactic violations in deep sedation (Heinke et al., 2004; Koe- et al., 2005; Wertz et al., 1998), epilepsy (Boatman et al., 2008; Lia- lsch et al., 2006), indicating that this kind of complex analysis sis et al., 2006), velo-cardio-facial (DiGeorge) syndrome (Baker following the rules of Western music can occur even in deep et al., 2005), Down’s syndrome (Lalo et al., 2005), children with sedation. certain forms of cleft palate and those with the Catch-22 syndrome Very recently, it was found by Boly et al. (2011) that in the veg- (Ceponiene et al., 2002) and, of course, in Alzheimer’s disease etative state, feed-forward processes are preserved whereas top- (Pekkonen et al., 1994; vMMN: Tales and Butler, 2006; Tales down processes are impaired. The effective connectivity was et al., 2008), dementia (Schroeder et al., 1995), intellectual disabil- measured by using the MMN for tones presented in a roving para- ity (Nakagawa et al., 2002; Ikeda et al., 2004) and mild cognitive digm (Baldeweg et al., 2004). The authors found that the only dif- impairment (MCI) (vMMN: Tales and Butler, 2006; Tales et al., ference between patients in the vegetative state and controls was 2008). an impairment of backward connectivity from frontal to temporal MMN-amplitude attenuation (at least for frequency change) can cortices, suggesting that a selective disruption of top-down pro- also index cognitive and functional degeneration in patients with cesses from high levels of a cortical hierarchy can lead to loss of an advanced stage of diabetes mellitus (Vanhanen et al., 1996), consciousness in brain-damaged patients, and can clearly differen- Parkinson’s disease (Karayanidis et al., 1995; Pekkonen et al., tiate the vegetative state from minimally conscious state. In addi- 1995b; Brønnick et al., 2010), HIV (Schroeder et al., 1994) and also tion to its neuroscientific relevance, the present approach could, in normal ageing (Gaeta et al., 2001; Kisley et al., 2005; Cooper according to Boly et al. (2011), constitute a new diagnostic tool et al., 2006). In addition, the MMN amplitude indexes decreased to quantify the level of consciousness electrophysiologically at cognition in different subnormal states such as sleep deprivation the patients’ bedside. (Raz et al., 2001), rapid eye movement (REM)-sleep deprivation The MMN can also reflect effects of hypnosis (Jamieson et al., (Zerouali et al., 2010) and in patients in the comatose (Fischer 2005; Kallio et al., 1999) and concentrative meditation (Srinivasan et al., 1999, 2006b) or PVS (Wijnen et al., 2007) state. and Baijal, 2007). Recently, Bisiachi et al. (2009) found smaller-amplitude MMN responses to tone-frequency change in newborns of less than 30 gestational weeks than in those with a longer gestational period. 11. The MMN as an index of the progression/severity of the This amplitude strongly correlated with several maturational fac- illness (Table 10) tors (gestational age, weight, length and cranial circumference). These results suggest, according to the authors, the role of the ges- In patients with schizophrenia, the MMN amplitude in particu- tational age as the main factor explaining cortical functioning at lar to frequency change is attenuated concomitantly with disease the early age. progression (for a meta-analysis, see Umbricht and Krljes, 2005). Consistent with this, several studies (Salisbury et al., 2002, 2007; 10. The MMN as an index of the level of consciousness (Table 9) Umbricht et al., 2006; Devrim-Ücok et al., 2008; Todd et al., 2008) showed that the frequency MMN deficit was more robust In patients in the comatose state, no MMN can usually be re- in chronic patients than in first-episode patients in whom the ef- corded unless a latent recovery process has started, leading to fect was smaller in size or had not yet emerged. In addition, Shino- the return of consciousness and cognitive capacities in the near fu- zaki et al. (2002) found that a temporary recovery of acute ture. Therefore, the MMN can be used as a tool in coma-outcome symptoms of schizophrenia patients was accompanied by an prediction (Kane et al., 1993, 1996; Fischer et al., 1999, 2004, MMN amplitude increase for tone-frequency change recorded at 2006a,b; Luauté et al., 2005; Morlet et al., 2000; Fischer and mastoids as polarity-reversed MMN (whereas the frontally re- Luauté, 2005; Daltrozzo et al., 2007; Vanhaudenhuyse et al., corded MMN did not correlate with this temporary recovery). In 2008). Moreover, the MMN peak latency measured at hospitalisa- addition, Urban et al. (2008), using the vMMN, found that the tion of survivors of coma caused by a severe head injury predicted, reduction of its amplitude for motion-direction deviants was larger to some extent, their expressive language ability and visuo-spatial for a more severe state and longer duration of the illness of the performance 1 year after the injury (Kane et al., 2000). Kotchoubey patient. (2007) stressed that in using the MMN for the objective assessment Moreover, MMN data also reflected clinical severity in patients of the patient’s remaining cognitive capacity, it is better to use with persistent developmental stuttering (Corbera et al., 2005), complex rather than simple sound stimuli which may result in se- dyslexia (Baldeweg et al., 1999; Stoodley et al., 2006), PTSD (Men- vere underestimation of the remaining capacity. For a meta-analy- ning et al., 2008) and in newborns and 6-month-old infants with sis of studies on the MMN in coma, see Daltrozzo et al. (2007). cleft palate (with clinical severity being determined as the ante- Recently, it was concluded by Vanhaudenhuyse et al. (2008) that rior–posterior extent of the cleft; Ceponiene et al., 1999, 2000a,b, MMN results in comatose patients converge to the conclusion that 2002). In a similar vein, Moberget et al. (2008) found that patients MMN is a very good predictor of recovery from coma, in particular exhibiting the most severe symptoms of ataxia, a sign of cerebellar in anoxic coma. cortical atrophy, produced smaller-amplitude MMN responses to In patients in the PVS, the frequency-MMN recovery occurred in tone-duration change than other patients. parallel with the recovery of consciousness (Wijnen et al., 2007; Studying stroke patients, Särkämö et al. (2008, 2009, 2010a, ad- Kotchoubey, 2007; Kotchoubey, 2007), showing a step-wise ampli- dressed amusia caused by a left or right middle cerebral artery R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 439

Table 11 The MMN predicts:

– Recovery from coma (Daltrozzo et al., 2007; Fischer and Luauté, 2005; Fischer et al., 1999, 2002, 2004, 2006a,b; Morlet et al., 2000; Naccache et al., 2005; Vanhaudenhuyse et al., 2008 (with the MMN being the best predictor of positive outcome, i.e., recovery without getting into the persistent vegetative state: Fischer and Luauté, 2005; Luauté et al., 2005); for a meta analysis, see Daltrozzo et al., 2007) – Recovery from the persistent vegetative state (Kotchoubey, 2007; Kotchoubey et al., 2007; Wijnen et al., 2007 (with the MMN recorded at hospilization also predicting the level of consciousness at discharge and even the extent of cognitive recovery during the next 2 years; Wijnen et al., 2007)) – Recovery from locked-in state (Ragazzoni et al., 2000) – Recovery from alcoholism (Kathmann et al., 1995; Pekkonen et al., 1998) – Social-skills learning in schizophrenia Kawakubo et al., 2007) – Social cognition in schizophrenia (Wynn et al., 2010) – Schizophrenia (ﬁrst-episode onset) risk (Bodatsch et al., 2011; Stone et al., 2010) – Progression of schizophrenia (Bodatsch et al., 2011; Devrim-Ücok et al., 2008; Kawakubo et al., 2007; Kiang et al., 2009; Light and Braff, 2005b; Salisbury et al., 2007; Umbricht and Krljes, 2005; Umbricht et al., 2006) – Dyslexia (Blitz et al., 2007; Leppänen et al., 2010; van Leeuwen et al., 2006; Stoodley et al., 2006) – Language development (Choudhury and Benasich, 2011; for a review, see Stix, 2011) – Language development in prematurely born children (Jansson-Verkasalo et al., 2003b, 2004, 2010) Fig. 8. Grand-average difference waves showing the duration mismatch negativity – Drug-therapy outcome in schizophrenia (Horton et al., 2010; Javitt et al., in the frontal (upper row) and central electrodes (lower row) for converting (dashed 2008; Lavoie et al., 2007; Schall et al., 1999) line) and nonconverting subjects (solid line). AR-C = at risk – coinversion, AR- – Working-memory performance decrement in chronic alcoholism (Ahveninen NC = at risk – no conversion (From Bodatsch et al., 2011). et al., 1999)

Table 12 The MMN shows increased genetic risk of: stroke. They found that amusia caused by a right-hemisphere damage, especially that to temporal and frontal areas, was more severe – Alcoholism (Rodriguez et al., 1998; Zhang et al., 2001) than that caused by a left-hemisphere damage. Furthermore, in – Asperger syndrome (Jansson-Verkasalo et al., 2005; Korpilahti et al., 2007) – Dyslexia (Blitz et al., 2007; Friedrich et al., 2004; Leppänen et al., 2002, amusic right-hemisphere patients, the severity of amusia corre- 2010; Leppänen and Lyytinen, 1997; Lyytinen et al., 2001, 2004, 2006; lated with weaker frequency-MMNm responses. Additionally, Maurer et al., 2009; Pihko et al., 1999; Richardson et al., 2003; van within the right-hemisphere-damaged group, the amusic patients Leeuwen et al., 2006) who had damage in the auditory cortex showed worse recovery – Language learning impairment (LLI) (Benasich et al., 2006; Choudhury and from amusia as well as weaker MMNm responses throughout the Benasich, 2011; Friedrich et al., 2004; Weber et al., 2005) – Schizophrenia in children (Schreiber et al., 1992), adults (Ahveninen et al., 6-month follow-up than did the non-amusic patients or the amusic 2006; Bodatsch et al., 2011; Brockhaus-Dumke et al., 2005; Hall et al., patients with no auditory-cortex damage. Also, the amusic patients 2007, 2009; Jessen et al., 2001; Magno et al., 2008; Michie et al., 2002; (both with and without auditory-cortex damage) performed worse Shin et al., 2009), and in Individuals with the CATCH syndrome (with a 108/158 108/ than did the non-amusic patients on a number of tests of different higher risk in individuals carrying the COMT Val than COMT 158Met allele) (Baker et al., 2005) aspects of cognitive performance (including working memory, – Specific language impairment (SLI) (Barry et al., 2008; Benasich and Tallal, attention and cognitive flexibility). These findings suggested, 2002; Benasich et al., 2006; Friedrich et al., 2004; Weber et al., 2005) according to the authors, domain-general cognitive deficits as the primary mechanism underlying amusia without auditory-cortex damage, whereas amusia with auditory-cortex damage is associ- Furthermore, in patients with schizophrenia, MMN data can ated with both auditory and cognitive deficits. predict future developments of the patient’s state and the effi- In conclusion, because of the large number of different disor- ciency of different treatments. For example, Umbricht et al. ders affecting the MMN, it appears that the MMN provides a gen- (2006) found that MMN deficiency at illness onset may index the eric illness measure, useful for assessing illness severity, as presence of a more pervasive brain pathology and be observed only suggested by Banati and Hickie (2009) for both the MMN and in a subgroup of patients, possibly in those with more precursors of microglia activation. schizophrenia or with an elevated risk of developing chronic schizophrenia. In addition, Kawakubo et al. (2007) found that the MMN in patients with schizophrenia was predictive of the acquisi- 12. The MMN in predicting illness course/drug response tion of social skills during a 3-month social skills training (prognosis) (Table 11) programme. Importantly, it was recently found by Lavoie et al. (2007) that A striking finding was that the recovery of the MMN in a coma- the deficient MMN amplitude for frequency change in patients tose patient strongly predicted, as already mentioned, the recovery with schizophrenia was corrected towards normal levels by a glu- of consciousness in the near future (Kane et al., 1993, 1996; Fischer tathione precursor, N-acetyl-cysteine (NAC), which improved their et al., 2000, 2006a,b; see also Fischer et al., 2008; Luauté et al., NMDAR function, supporting the role of the NMDARs in MMN 2005; Vanhaudenhuyse et al., 2008; for a meta-analysis, see Dal- generation. trozzo et al., 2007). In addition, the MMN in patients in the PVS re- The MMN improvement was, however, found in the absence of corded at hospitalisation predicted the magnitude of cognitive any robust changes in clinical-severity indices, even though they recovery even 2 years ahead (Wijnen et al., 2007; see also Kotchou- were observed in a larger and more prolonged clinical study (La- bey, 2007; Kotchoubey et al., 2007). voie et al., 2007; Berk et al., 2008a,b; for supporting evidence, 440 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458

Table 13 MMN indexes central-auditory-processing improvement in:

– Autistic patients (with frequency processing better than in Controls; Gomot et al., 2002) – Children with speciﬁc language impairment (SLI) receiving language and speech training (Pihko et al., 2007) – Cochlear-implant (CI) patients as a function of time since CI installation (Kraus et al., 1993b; Lonka et al., 2004; Ponton et al., 2000; Ponton and Eggermont, 2001) and training (Ponton et al., 2009) – Dyslexic children presented with consonant–vowel syllables with lengthened formant-transition duration (Bradlow et al., 1999) – Dyslexic children receiving audio-visual rehabilitation (Kujala et al., 2001) – Early Child development, mainly speech-sound learning (Alho and Cheour-Luhtanen, 1997; Alho et al., 1990; Carral et al., 2005; Cheour et al., 1998b, 2002a,b; Cheour- Luhtanen et al., 1996; Choudhury and Benasich, 2011; Draganova et al., 2005, 2007; Gomes et al., 2000; Gomot et al., 2000; Fellman and Huotilainen, 2006; Huotilainen et al. 2005, 2007; Kraus and Cheour, 2000; Kraus et al., 1992, 1993a, 1999; Kuhl, 2004; Kujala et al., 2004; Kushnerenko et al., 2002, 2007; Leppänen et al., 2004; Lovio et al., 2009; Lyytinen et al., 2001, 2006; Molholm et al., 2004; Novitski et al., 2007; Oades et al., 1996; Pang et al., 1998; Pihko et al., 2005; Tanaka et al., 2001; Trainor et al., 2001, 2003; Uwer and von Suchodoletz, 2000) – Early-onset blindness (Kujala et al., 1995a,b, 2000b, 2005b; Alho et al., 1993) – Epileptic patients with vagus-nerve stimulation (Borghetti et al., 2007) – Infants operated for craniosynosthosis (Huotilainen et al., 2008) – Patients with hearing aids (Rudner et al., 2008) – Patients with ultrasound hearing aids (Hosoi et al., 1998) – Schizophrenic patients progressing from the Acute to Post-Acute Phase (Shinozaki et al., 2002) – Sleeping infants exposed to auditory discrimination training (Cheour et al., 2002a) – Stroke patients as a function of time since stroke onset (Ilvonen et al., 2003; Särkämö et al., 2010a,) – Stroke patients receiving music or audio-book stimulation (Särkämö et al., 2010a,)

see Coyle, 2006; Schwieler et al., 2008). Hence, an MMN enhance- discriminate non-native phonemes was preserved in prematurely ment may precede improvement in clinical severity, which, born infants. Moreover, it was also found that the better this ability according to the authors, highlights the possible utility of the was preserved (reflected by larger MMN-amplitudes for non-na- MMN as a biomarker of treatment efficacy, and, therefore, also as tive contrasts) at the age of 1 year, the worse was the performance a tool in drug development (see also Javitt, 2007, 2009; Javitt in native-language tests at the age of 2 years. Thus, decline in sen- et al., 2008). Previously, in the Schall et al. (1999) study, the sitivity to non-native phonemes, as reflected by MMN attenuation MMN predicted the outcome of clozapine therapy in patients with for non-native phoneme contrasts, served as a positive predictor schizophrenia. Moreover, more recently, Horton et al. (2011) found for further age-appropriate language development. that clozapine increased the MMN amplitude for tone-frequency Moreover, the Weber et al. (2005) MMN data showing a deficit change, with increasing doses being associated with increasing fre- of prosodic perception in 5-month-old infants suggested that the quency-MMN amplitudes (but attenuated that for tone-duration MMN might be taken as an early marker of SLI risk. increments and decrements). Very recently, it was found by Bodatsch et al. (2011) that in individuals with a high schizophrenia risk, the MMN for tone-dura- 13. The MMN as an index of genetic disposition to different tion decrement (but not for tone-frequency change) was lower in pathologies (Table 12) amplitude (resembling that of first-episode patients) in those patients who subsequently, during an observation period of 2 years, Some studies suggest that in schizophrenia there is a genetic converted to the first-episode psychosis than that in those who contribution to its aetiology. An MMN-amplitude attenuation for did not convert to psychosis during that period (Fig. 8). This result duration increment was found by Michie et al. (2002) and that was remarkable as these groups were thought to share a similar for frequency change by Jessen et al. (2001) in symptom-free clinical risk of psychosis at baseline. The authors proposed that first-order relatives of patients with schizophrenia but studies with the duration-MMN reduction in their converters might reflect both larger samples have either not confirmed these findings (Bramon structural and neurochemical alterations associated with the con- et al., 2004) or showed a trend only (Price et al., 2006). However, version to psychosis. Furthermore, such results do not, according to Hall et al. (2006) recently found a significant, although modest, ge- the authors, only contribute to the prediction of conversion and netic correlation between the duration-increment MMN-ampli- lead time to it but also contribute to a more individualised risk tude deficit and schizophrenia, and therefore suggested that the assessment and hence also to risk-adapted prevention. In addition, MMN is a potentially valid endophenotype in schizophrenia. By recently, Stone et al. (2010) found in subjects at risk of psychosis, contrast, Ahveninen et al. (2006), using the MMN to tone-fre- an attenuated frontal-MMN amplitude for tone-duration incre- quency changes in their study of twin pairs discordant for schizo- ment and, further, that this MMN deficit was related to reduced phrenia, obtained results suggesting, according to the authors, that thalamic glutamate plus glutamine and NMDA levels. MMN abnormalities in patients with schizophrenia might reflect Further, a phonological deficit in young elementary-school chil- predominantly state-dependent neurodegeneration. A similar con- dren, reflected by a reduced MMN amplitude to syllable change clusion was made by Magno et al. (2008) on the basis of their dura- (but not that to simple frequency change), appeared to predict tion- and frequency-MMN amplitude results. their dyslexia risk (Blitz et al., 2007). Also, in very prematurely In patients with the 22q11 deletion syndrome, the considerably born children, the MMN amplitude for consonant change at 4 years increased rates of schizophrenia are attributed to the loss of one al- of age predicted, as already mentioned, naming performance at the lele containing the catechol-O-methyl transferase (COMT) gene. age of 6 years (Jansson-Verkasalo et al., 2003b, 2004). Furthermore, Symptom-free carriers showed attenuated MMN amplitudes for Jansson-Verkasalo et al. (2010) found that the perceptual narrow- both simple auditory (duration and frequency) and syllabic ing for non-native phoneme contrasts occurring in normal healthy changes as well as corresponding decrements in behavioural sound infants from 6 to 12 months of age (Kuhl et al., 1991; Kuhl, 2004), discrimination (Baker et al., 2005). Moreover, these MMN and reflected by attenuated MMNs for non-native phoneme contrasts behavioural effects were stronger in subjects carrying the COMT in these infants (Cheour et al., 1998b), does not occur in very pre- 108/158Met allele on their single intact chromosome (associated maturely born infants who showed large MMNs even for non-na- with a greater risk of schizophrenia) than in those carrying the tive phoneme contrasts. This indicates that the ability to COMT 108/158Val allele (a smaller schizophrenia risk). The authors R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 441 concluded that because the MMN depends on the NMDAR function cooperate. The MMNs in the first two sessions were quite small (Javitt et al., 1996), the MMN modification by the COMT Val 108/ in amplitude for sound-duration and -frequency changes, but be- 158Met polymorphism points towards abnormal dopamine–gluta- came larger thereafter, disclosing normal-like amplitudes at mate interactions in the MMN-generator system. 3 months post-stroke (Fig. 3). Furthermore, the duration-MMN Furthermore, children with a parent or sibling with dyslexia amplitude increase correlated with the improvement in the Boston have an increased risk of dyslexia, and this is reflected by their Diagnostic Aphasia Examination test scores from 10 days to attenuated MMN amplitudes to simple frequency (Maurer et al., 3 months. 2003) or duration changes (Friedrich et al., 2004; Leppänen et al., Subsequently, as already mentioned, it was found by Särkämö 2002) or to phoneme-category change (van Leeuwen et al., et al. (2010b) that the recovery of the MMNm and behavioural dis- 2006). Similarly predictive were the lack of a left-hemisphere crimination can be expedited with music and audio-book stimula- MMN response to speech-sound change (Maurer et al., 2003) and tion. Dividing their 60 middle cerebral artery stroke patients into the bilateral distribution of this MMN response (Maurer et al., music and audio-book listening and control groups, these authors 2009). recorded patients’ MMNm to frequency and duration changes in Also, studying 6-month-old infants with a family history of lan- a binaurally delivered complex tone. (In addition, all patients re- guage-related learning difficulties, Benasich et al. (2006) found ceived the normal hospital treatment and rehabilitation of stroke that these infants had smaller mismatch responses (MMRs; the po- patients, of course.) It was found, as expected, that the MMNm sitive-polarity analogue of MMN; Maurer et al., 2003) than those of amplitudes were, in general, smaller in the lesioned than opposite control infants with no such family history. In addition, as already hemisphere. Importantly, the frequency-MMNm amplitude in- mentioned, Barry et al. (2008) obtained MMN data suggesting that creased more in both music and audio-book groups than in the parents of children with SLI have a shorter auditory sensory-mem- control group during the 6-month post-stroke period. In addition, ory duration than that of control parents. the duration-MMNm amplitude increased more in the audio-book Very recently, a major breakthrough in the research of the ge- group than in the other groups. Moreover, the frequency-MMNm netic background of dyslexia was made by Roeske et al. (2011) amplitude changes correlated with the behavioural improvement who found that a certain marker (rs4234898), located on chromo- of verbal memory and focussed attention induced by music listen- some 4q32.1, was associated with the MMN deficit for a CV-sylla- ing. The authors interpreted their results as demonstrating that the ble change. Moreover, this MMN was also associated with a mere listening to music or speech starting soon post-stroke onset two-marker haplotype of rs4234898 and rs11100040, one of its can induce long-term plastic changes in early sensory processing, neighbouring single-nucleotide polymorphisms. The authors con- as reflected by the MMNm enhancement, which, in turn, may facil- cluded that their results suggest a possible transregulation effect itate the recovery of higher cognitive functions. These remarkable on SLC2A3, the predominant facilitative glucose transporter in results encourage the use of music and speech stimulation in the neurons, which might lead to glucose deficits in dyslexic children rehabilitation of stroke patients in particular in the early post- and could explain their attenuated MMN in passive listening tasks, stroke period. as observed in this study. Moreover, in a very recent study of the The deficit of the auditory/language system may be inborn, as is same group, Czamara et al. (2011) obtained results pointing to the case in developmental dyslexia, SLI or in the autism spectrum. an association between the late MMN for speech-sound change Because of the high prevalence of these disorders and their devas- and rare variants in a candidate region for dyslexia. tating effects on the individual’s academic success and thereby self-esteem, early identification and remediation methods should be developed. Recent evidence fortunately shows that the aberrant 14. The MMN as an index of central auditory processing neural substrate of language deficits can be ameliorated with improvement or recovery (Table 13) appropriate intervention. For instance, Pihko et al. (2007) found that language-impaired children at the age of 5–6 years benefited Spontaneous and training-induced improvement. A number of from language and speech exercises. Furthermore, this training studies in normal healthy subjects (Atienza and Cantero, 2001; also changed the neural processing of speech sounds so that the Kraus et al., 1995; Gottselig et al., 2004; Menning et al., 2000; MMNm for syllable changes became stronger in the left Tremblay et al., 1997, 1998; Näätänen et al., 1993; for a review, hemisphere. see Kujala and Näätänen, 2010) demonstrated that the MMN re- In dyslexic first-grade children, intervention, judging from an flects plastic neural changes associated with discrimination learn- MMN-amplitude enhancement, induced neural plastic changes ing. Therefore, it is a very attractive tool for following up recovery and ameliorated reading difficulties (Kujala et al., 2001). These 7- and intervention effects in various patient groups, for instance, in year-old school children were exposed to an audiovisual training stroke, coma, PVS or cochlear-implant (CI) patients or in children programme with no linguistic items (Audilex Program; Karma, with developmental disorders. 1999) for 7 weeks. Their reading skills as well as the MMN elicited In stroke and other brain-injured patients, it would be impor- by tone-order reversals were assessed before and after this training tant to determine the changes in perceptual abilities very soon period. After this period, the training group made fewer reading er- after the damage. However, such patients are often unable to coop- rors and tended to read faster than did the age-matched control erate or to understand instructions, particularly if the neural net- group of dyslexic children. Moreover, the MMN amplitude for work involved in speech perception is affected. Aaltonen et al. tone-order reversals was increased in the training group but not (1993), in an early study, found that the MMN to speech-sound in the control group. These results consequently showed that read- change was abolished in posterior left-hemispheric stroke patients, ing deficits can be ameliorated with audiovisual training and, fur- whereas an anterior left-hemispheric stroke had no such effect. By ther, that the improvement of reading skills is associated with contrast, the tone-frequency MMN was normal in both patient plastic neural changes of the central auditory system as reflected groups. Subsequently, as already mentioned, Ilvonen et al. (2003) by the MMN enhancement. Furthermore, these positive training followed up the recovery of left-hemisphere stroke patients, results were obtained by using no linguistic stimuli, which sug- recording the MMN at 4 and 10 days, and then at 3 and 6 months gests that impaired auditory-perceptual processes other than defi- after stroke onset. Language-comprehension tests (Boston Diag- cient phonology may also contribute to reading impairments. nostic Aphasia Examination and Token tests) were also adminis- Similar effects were also obtained with extremely low-birth- tered in all but the first session in which the patients could not weight children of 6 years of age (Huotilainen et al., 2011). These 442 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 children were allocated to a 5-week training programme with comparable response. Furthermore, Kelly et al. (2005) reported a Audilex (training group) or to a control group playing computer similar failure to obtain the MMN from CI users defined as poor games. Before and after the intervention, auditory ERPs to sound performers. In addition, Lonka et al. (2004), in longitudinally track- changes were recorded and reading-related skills were assessed. ing the changes in the language function after a small group of deaf The MMN responses to frequency and duration deviants increased adults received a CI, reported that while the MMN was absent for in amplitude in the Audilex training group but not in the control- approximately 12 months post-surgery, the response subsequently game group. However, the reading skills were similar in the two emerged, first for a large vowel contrast, and then later for more groups after invention and 2 years later. subtle vowel distinctions. Converging behavioural improvement As already mentioned, the shortening of the abnormally long in a speech-discrimination test was also observed during the fol- MMN-peak latency and the MMN-amplitude recovery in drug- low-up period. resistant adult epilepsy patients after vagus-nerve stimulator Trautwein et al. (1998) and Ponton et al. (2000), who assessed implantation was recently found by Borghetti et al. (2007), sug- the CI users with non-speech stimuli, obtained results that relate gesting a positive effect of vagus-nerve stimulation on central back to both speech performance and training. First, Trautwein auditory processing in these patients. et al. (1998) found that while psychophysical thresholds were Drug-induced improvement.InTable 1, short-term effects of dif- superior to MMN-estimated discrimination thresholds for duration ferent drugs are reviewed. In addition, in the foregoing, some contrasts in the control group, the opposite was found for a group favourable drug effects in schizophrenia patients were already of adult CI users. Thus, in several individuals, there was neuro- mentioned (NAC, Lavoie et al., 2007; clozapine, Schall et al., physiological evidence of discrimination based on the MMN, with- 1999; Horton et al., 2011). Moreover, very recently, Dulude et al. out the individual being able to behaviourally perceive the (2010) found that nicotine, relative to placebo, not only prolonged difference. Based on the apparent separation of discrimination at the peak latency of the duration MMN but also increased its ampli- the level of sensation versus perception, Ponton et al. (2009) devel- tude in patients to a level comparable with that seen in controls. By oped a protocol based on pre-training MMN evaluation to deter- contrast, the frequency MMN was not affected. The authors con- mine whether there was neurophysiological evidence of cluded that these preliminary findings demonstrate that acute nic- discrimination in the absence of perceptual discrimination. This otine can normalise temporal aspects of central auditory knowledge was then used to develop a training protocol which re- processing in patients with schizophrenia, an effect that might be sulted in significant improvements in frequency discrimination as mediated by the activation of a7 nicotinic acetylcholine receptors, well as combined consonant and vowel discrimination in a single the functioning of which is diminished in schizophrenia. The highly trained adult implant user. authors further concluded that these ameliorating effects of nico- As noted by Roman et al. (2005), there is a clear application of tine may have implications for understanding the increased tobac- the MMN in clinical testing and training of CI populations. How- co smoking in schizophrenic patients and for developing nicotinic ever, refinements are needed in methods and recording technique, pharmacotherapeutics to alleviate sensory-memory impairments to optimise the protocols for frequency discrimination and speech in schizophrenia. In addition, a positive effect of nicotine in both testing. smoking and non-smoking normal healthy subjects on temporal- Recently, music perception of CI users was also studied by using interval processing (indexed by an MMN-amplitude enhancement the MMN. Koelsch et al. (2004) found that CI users can perceive to ISI deviants) was recently found by Martin et al. (2009). timbre, judging from the MMN elicited by timbre deviants in them Furthermore, very recently, it was found by Sawada et al. (2010) (the timbre MMN: Tervaniemi et al., 1997; Toiviainen et al., 1998). that in children of 7–13 years of age with attention deficit hyperac- Moreover, music-syntactic violations elicited an early right ante- tivity disorder (ADHD), the intake of osmotic-release methylpheni- rior negativity which might be interpreted as a music-syntactic date (MPH) increased the MMN amplitude for tone-frequency MMN in CI users, which again was considerably smaller in ampli- change and simultaneously alleviated the severity of their ADHD tude than that in controls but nevertheless showed that CI users’ symptoms. central neural mechanisms underlying music perception are acti- Central auditory discrimination and training in CI users. For pro- vated by musical stimulation (even though they reported that they foundly deaf individuals, a sensation of hearing can be restored had difficulties in discriminating musical information and that by a CI. CIs are currently the most successful and widely used sen- they were in general frustrated by their inability to accurately hear sory neuroprotheses. In a large percentage of those individuals fit- music). Consistent with this, subsequently, Sandmann et al. (2010), ted with a CI, a remarkable degree of language communication via by using the multi-feature MMN paradigm with six different levels the auditory domain is restored. Nevertheless, objective, non- of the magnitude of deviation for each feature (Pakarinen et al., behavioural measures of auditory discrimination such as the 2007)(Fig. 1) found that CI users had difficulties in discriminating MMN remain a potentially powerful, but as yet under-utilised, small changes in the acoustic properties of musical sounds. It was application for this population. concluded by Sandmann et al. (2010) that this type of paradigm The first MMN recordings from CI users were reported by Kraus could be of substantial clinical value by providing a comprehensive et al. (1993b) who employed speech-syllable contrasts. For the profile of the extent of restored hearing in CI users. Moreover, an group of individuals investigated in that study, Kraus et al. found inverse relationship was found between the MMN amplitude and that the MMN elicited from CI users was remarkably similar to that the duration of profound deafness; however, there was also evi- obtained from a group of normal-hearing listeners. These results dence for experience-related plastic changes as a function of the suggested that despite the differences in peripheral input, the brain duration of CI use (Sandmann et al., 2010). was processing basic speech units in a very similar fashion. Ultrasonic hearing. Some profoundly deaf individuals can per- Ponton et al. (2000; see also Ponton and Don, 1995, 2003; Pon- ceive ultrasound only, which occurs by bone conduction. Hosoi ton and Eggermont, 2007), recording the MMN evoked by a single et al. (1998) found that in normal healthy subjects, ultrasound contrast in stimulus duration from groups of normal-hearing modulated by different speech sounds produces discriminable adults and adult CI users, also found that MMNs of the two groups stimuli, which points to the possibility of developing ultrasonic were quite similar. In a more detailed study in which subjects were hearing aids as an alternative for CIs for those profoundly deaf indi- separated into groups based on benefit gained by using a CI, Groe- viduals who can sense ultrasound only. Hosoi and colleagues deliv- nen et al. (1996) reported that whereas the group classified as good ered ultrasound to the right sterno-cleidomastoid through a performers showed an MMN, poor performers failed to generate a ceramic vibrator. Substantial N1m responses were elicited by both R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458 443 audible air-conducted sound stimuli and bone-conducted ultra- development, for example, the prediction of coma or PVS outcome sonic stimuli. Further, when the (40 kHz) ultrasound stimuli mod- and even that of the extent of the cognitive recovery of these pa- ulated by vowels/i/ and/a/ were presented as standards and tients (during the next 2 years; Wijnen et al., 2007), or the moni- deviants, respectively, substantial MMNm responses were ob- toring of, and predicting, the progress of schizophrenia. served for deviants, indicating that the two sounds were discrimi- Moreover, importantly, very recent results of Bodatsch et al. nable. Moreover, the equivalent current dipole (ECD) for the (2011) showed that the duration MMN can predict, in individuals MMNm was located within the auditory cortex, in the vicinity of in schizophrenia-risk group, conversion to first-episode schizo- the MMNm observed in a separate condition with the respective phrenia and even when this would occur in the period of 2 years. air-conducted stimuli. The authors also tested 53 profoundly deaf In addition, another recent study (Banati and Hickie, 2009) sug- individuals behaviourally and found that 28 of them could detect gested that the duration and frequency MMNs might enable one ultrasonic stimuli and, further, that 11 of them could discriminate to online monitor local disease activity and neuronal death in pa- ultrasounds modulated by different speech signals. These results tients with schizophrenia. provide, according to the authors, the rationale for developing Furthermore, in case of schizophrenia (see Lavoie et al., 2007; ultrasound hearing aids, which do not require surgery, for those Horton et al., 2011; Javitt et al., 2008; Banati and Hickie, 2009; profoundly deaf who can sense ultrasound. see also Berk et al., 2008a,b), and perhaps of other severe neuro- Improved discrimination in early blind individuals. A special case psychiatric, neurological and neurodevelopmental disorders, too, of enhanced central auditory function of compensatory nature the MMN can be used as an online index of treatment efficacy. can be encountered in early blinds. The MMN elicited by sound- As far as dyslexia is concerned, such a use of the MMN in assessing location changes was larger in amplitude in early-blind individuals the effectiveness of different rehabilitation programmes is already than in sighted control subjects (Kujala et al., 1995a). (The MMN well established (Kujala et al., 2001). In addition, in the foregoing, was recorded while subjects’ attention was directed to the tactile we reviewed results of Särkämö et al. (2010a) demonstrating how modality.) This suggests enhanced spatial auditory perception the MMNm can be used in evaluating the effectiveness of different and discrimination in the blind. The result is consistent with auditory stimulus material in an attempt at facilitating stroke behavioural data showing that the blind are more accurate than recovery. the sighted in detecting loci of origin of sounds (Lessard et al., Some further clinically important uses of MMN data are, among 1998; Röder et al., 1999). The scalp topography of MMN response other things, the possibility to determine the dyslexia risk in new- of Kujala et al., 1995a) did not differ between the two groups, how- borns and infants (Maurer et al., 2003; Leppänen et al., 2002; van ever. It, therefore, behaved differently from the N2b (Näätänen Leeuwen et al., 2006; Friedrich et al., 2004), the magnitude of cog- et al., 1982; for a review, see Näätänen and Gaillard, 1983) re- nitive and functional deficit associated with different clinical con- sponse elicited by auditory targets in an active discrimination conditions (Näätänen et al., 2011a) and the degree of severity of dition, which had a scalp topography in the blind that was different conditions (in Schizophrenia: Umbricht et al., 2006; Dev- posterior to that in the sighted (Kujala et al., 1992, 1995a,b). Thus, rim-Ücok et al., 2008; Todd et al., 2008; Light and Braff, 2005a,b; the MMN appears to be hard-wired in the central auditory system Salisbury et al., 2002, 2007; Stuttering: Corbera et al., 2005; PTSD: of the blind, reflecting plastic changes within the auditory cortex, Menning et al., 2008; Cleft Palate: Ceponiene et al., 1999, 2000, whereas the generators of the N2b, associated with attentional 2002; Stroke and Aphasia: Ilvonen et al., 2003; Pettigrew et al., processes in audition, may become part of the neural system nor- 2005; Särkämö et al., 2010b; Dyslexia: Baldeweg et al., 1999; Kuj- mally subserving vision. This was supported by imaging studies ala et al., 2001). suggesting occipital activity in the blind during auditory change- One might wonder why auditory discrimination is affected in detection tasks (Kujala et al., 1995b, 2007, 2000b, 2005b). such a large number of different clinical conditions (Table 1), which indeed points to a possible common pathology shared by these different conditions. A major part of this common pathology 15. Concluding discussion appears to be found in the deficient functioning of the NMDAR system, as there is very strong evidence indicating that MMN defi- This article has shown that central auditory processing, as in- ciency reflects deficient NMDAR functioning (Javitt et al., 1996; dexed by the MMN and the MMNm, is affected in a wide range Näätänen et al., 2011a). For example, sub-anaesthetic doses of ket- of different clinical conditions and in ageing. Most of these effects amine, an NMDAR antagonist, attenuated the MMN amplitude of are seen as indexing decreased auditory discrimination accuracy. healthy volunteers (Kreitschmann-Andermahr et al., 2001: In some cases, however, the duration of auditory short-term sen- MMNm; Umbricht et al., 2000, 2002), monkey (Javitt et al., sory memory, essential, for instance, in speech perception and 1996), rat (Tikhonravov et al., 2008, 2010) and mouse (Ehrlichman understanding, is affected. This further decreases automatic dis- et al., 2008), and caused schizophrenic kinds of psychotic experi- crimination when SOAs are prolonged. Moreover, these data also ences in normal subjects (Umbricht et al., 2000, 2002; see also show effects on mechanisms controlling for involuntary shifts of Sauer and Volz, 2000; Pang and Fowler, 1999). Furthermore, keta- attention (passive attention) or on those determining the charac- mine disturbed the performance in a memory task in normal sub- teristics of backward masking in different patient groups. jects in a manner similar to that normally observed in Consequently, this article has shown that the MMN provides a schizophrenia, as shown by Umbricht et al. (2000). Importantly, unique window to the neuropsychology of central auditory pro- it was found by Umbricht et al. (2002) that the smaller was the cessing, and hence a possibility for the objective assessment of subject’s MMN amplitude for frequency and duration changes be- auditory discrimination and sensory memory, in different patient fore ketamine administration, the stronger were the psychotic groups, which previously could be mainly inferred from behav- experiences caused by the drug. Therefore, the authors concluded ioural performance only. This is a major improvement, in particular that the MMN in the normal state of consciousness indexes the when considering the special communicational and motivational vulnerability to disruption of the NMDAR system in normal healthy problems with patients, deducting from the reliability of the re- subjects. Consequently, the MMN appears to index the functional sults, and the fact that reliable behavioural measurements are state of the NMDAR-mediated neurotransmission even in subjects not at all possible in some clinical groups and often in small in- demonstrating no psychopathology and may therefore ‘‘become a fants. These new possibilities in particular involve the improved useful tool in studies on NMDAR functioning in humans’’ (Umb- monitoring of the patient’s state and the prediction of its future richt et al., 2002). 444 R. Näätänen et al. / Clinical Neurophysiology 123 (2012) 424–458

Consequently, the MMN attenuation observed in a large num- creases the amount of information that can be gathered within a ber of different clinical populations and conditions (Table 1) can, specific time period. Equally as important are methods that would to a large extent, be accounted for by NMDAR dysfunction associ- further reduce the amount of time required to acquire sufficient ated with different clinical conditions. For instance, as already re- data that reach some objective criteria of identification. viewed, the dysfunction of the NMDAR system plays a central One such approach has recently been described by Rahne et al. role in the brain pathology of schizophrenia and in particular in (2008). In this modification to standard averaging, individual the cognitive deficit in these patients (Javitt et al., 1996; Coyle, epochs to stimulus presentation are sorted on the signal-to-noise 2006). Moreover, the cognitive decline occurring in several other ratio (SNR) to each individual epoch. This epoch with the highest neurocognitive abnormalities might also be accounted for, at least SNR, that is, the lowest background noise, is added first to average. in part, by the dysfunctional NMDAR system (Näätänen et al., Individual epochs are then added to the average in rank order 2011a). Alzheimer’s disease, stroke, traumatic brain injury and based on the background noise in each epoch. Thus, averaging is even normal ageing are accompanied by widespread dysfunctions continued until the SNR is optimised. Objective or statistical detec- in the NMDAR system (Lipton and Rosenberg, 1994; Magnusson tion of these responses could then easily be achieved by using one et al., 2010; Biegon et al., 2004; Dalkara et al., 1996; Izquidero, of a number of online algorithms involving randomisation or boot- 1991; McGeer and McGeer, 2010). For example, Dhawan et al. strapping analyses such as those proposed by, for example, Ponton (2010) recently found that middle central artery occlusion in rat et al. (1997) or Fujioka et al. (2005). Moreover, further progress to resulted in widespread decreases in NMDAR density in brain re- this end has been very recently made by Cong et al., 2009, 2010, gions extending far beyond the infarct area, including even regions 2011); Burger et al. (2007); Kalyakin et al. (2007, 2008a,b, 2009) contralateral to the lesion important to cognitive function. Dhawan and Marco-Pallares et al. (2005). et al. (2010), therefore, concluded that a persistent and widespread Needless to say, a lot of work is still needed to bring the MMN loss of NMDARs may contribute to cognitive and other neurological methodology to clinics as a tool of everyday patient work in which deficits in stroke patients, which cannot be localised to the side of reliable measurements have to be carried out at the level of infarction. individual patients (Bishop and McArthur, 2004; Boly et al., Importantly, this NMDAR functional deficiency may hence ac- 2011; Taylor and Baldeweg, 2002; Lang et al., 1995), but studies count for, besides decreased discrimination accuracy, even the cog- as those reviewed in the foregoing have shown that this goal is nitive and functional decline occurring in a number of these both valuable and probably also attainable. conditions (Table 8) (for a review, see Näätänen et al., 2011a). It is, for instance, well established that the adequate functioning of Acknowledgements the NMDAR system plays a crucial role in the initiation of long- term and working memories (Javitt et al., 1996; Newcomer et al., RN and KK were supported by Grant # 8332 from the Estonian 1998). Therefore, the MMN and MMNm deficiency reflecting a defi- Science Foundation. TK was supported by Grant # 128840 and RN cient NMDAR function can also index cognition and its decrement by grant # 122745 from The Academy of Finland. CE was supported in ageing and in different neurological and neuropsychiatric abnor- by grants from ICREA Academia 2010 and CSD2007-00012, malities (Näätänen et al., 2011a). Consequently, as an easily acces- EUI2009-04806, PSI2009-08063, and SGR2009-11. We wish to sible non-invasive measure of the functional condition of the acknowledge Ms. Piiu Lehmus’ skilful and patient text-editing NMDAR system, the MMN (MMNm) might play an important role work. in future attempts at understanding what is common in the different neuropsychiatric diseases and abnormalities and what is spe- References cific to each of them (Umbricht et al., 2003). These conclusions are further supported by studies showing that pharmacological Aaltonen O, Tuomainen J, Laine M, Niemi P. Cortical differences in tonal versus manipulations enhancing cognition also enhance the MMN vowel processing as revealed by an ERP component called mismatch negativity. (MMNm) response (Baldeweg et al., 2006; Dunbar et al., 2007; Brain Lang 1993;44:139–52. Ahissar M. Dyslexia and the anchoring-deficit hypothesis. 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