Early Occurrence Of Auditory Change Detection In The Human Brain

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1 Early Occurrence Of Auditory Change Detection In The Human Brain SABINE GRIMM University of Leipzig, Germany (formerly University of Barcelona, Spain) ICON, Brisbane, July 28 th, 2014

Auditory deviance detection in animals and humans 2 The auditory system constantly scans the acoustic input for regularities and respectively for unexpected, novel events!

2 Auditory deviance detection in animals and humans 2 The auditory system constantly scans the acoustic input for regularities and respectively for unexpected, novel events! Human scalp potential Animal multi-/single-unit responses from A1 from: Schröger, 2005 from: Ulanovsky et al., 2004 Astonishing similarities in their characteristics! (see Nelken et al., 2007) similar relation to magnitude of change, to probability of deviant events, to the number of standards; similar local effects in 50-50% conditions;

, 2010) Release from SSA occurs already within the first 30 ms after deviant onset (Pérez- González et al.

3 Early occurrence of animal SSA 3 Animal studies Neurons that show SSA have been reported in primary auditory cortex (A1, Ulanovsky et al. 2003, 2004) and in subcortical areas (IC and MGB, Pérez-González et al., 2005; Antunes et al., 2010) Release from SSA occurs already within the first 30 ms after deviant onset (Pérez- González et al., 2005; von der Behrens et al., 2009) Human studies Generators of MMN are localized in auditory cortical areas and frontal cortex (Alho, 1995; Huotilainen et al., 1998) MMN peaks between ms

4 Are there any human correlates of deviance detection at earlier latencies than those of MMN, that could bridge the gap to novelty-related responses recorded in animal studies?

5 Is human deviance detection a multi-stage process? In principle, one should be able to use the Auditory Evoked Potential, in its whole complex morphology, to tackle this question. 1. The Auditory Brainstem Response [ABR].

6 Is human deviance detection a multi-stage process? In principle, one should be able to use the Auditory Evoked Potential, in its whole complex morphology, to tackle this question. 1. The Auditory Brainstem Response [ABR]. 2. The Middle Latency Response [MLR].

7 Is human deviance detection a multi-stage process? In principle, one should be able to use the Auditory Evoked Potential, in its whole complex morphology, to tackle this question. 1. The Auditory Brainstem Response [ABR]. 2. The Middle Latency Response [MLR]. 3. The Long-Latency Auditory Evoked Potential [including the MMN]. N1 MMN -1 µv 300 ms Típico Discrepante Onda diferencia

Nbm enhancement for 2580 frequency Hz 1770 Hz deviants 1200 Hz 800 Hz More anterior and medial activations (Heschl s c gyrus) for Nbm

8 Early EEG correlates of deviance detection 8 EEG: Nb enhancement for frequency deviants a Grimm et al. (2011). Psychophysiology. deviant 800 Hz 3730 Hz 2580 Hz 1770 Hz 1200 Hz 800 Hz deviant 3730 Hz b Nbm MEG: standard 800 Hz standard difference 3730 Hz 3730 Hz Nbm enhancement for 2580 frequency Hz 1770 Hz deviants 1200 Hz 800 Hz More anterior and medial activations (Heschl s c gyrus) for Nbm compared to control 800 Hz control 3730 Hz MMNm Recasens et al. (2012). Cer Cortex. [p=0.012] MMNm ~ 42 ms!! Tone duration: 50 ms SOA: 293 ms

9 Early EEG correlates of deviance detection 9 EEG: Nb enhancement for MF pitch changes -2 µv Alho, Grimm et al. (2012). EJN N ms Missing fundamental (MF) stimuli: µv frequency ms frequency 0.5

2006, Brain Res + deviant control Grimm et al., 2012, Biol Psychol Cornella et al.

10 Early EEG correlates of deviance detection 10 Na enhancement for location deviants - Na [24 ms] - Na [20ms] + Na difference topography Sonnadara et al. 2006, Brain Res + deviant control Grimm et al., 2012, Biol Psychol Cornella et al. 2012, PLoS One Pa/Nb enhancement for SOA shortenings Leung, Recasens, Grimm, Escera, 2013, Clin Neurophysiol

11 So far, so good 11 Deviance-related effects occur at multiple latencies including much shorter ones than those of the MMN!!! Early deviance effects occur at different components (Na, Nb: ms) of the Middle Latency Response depending on the stimulus feature: Some indication that the early effects reflect an enhanced deviance processing rather then repetition suppression (based on experiments controlling for stimulus probability) MEG: Nbm effect is localized near primary auditory cortex whereas MMNm shows more posterior and lateral activations Functional role of these multiple stages of deviance detection? * acting in a complementary, hierarchical, interactive, or a redundant manner?

12 Can the early deviance-related effects be observed in the context of more complex regularities?

13 Early detection of violation within complex regularities 13 FEATURE CONJUNCTION DEVIANTS Hz 45% 5% 1200 Hz 5% 45% Hz 45% 45% 1200 Hz 5% 5% Althen, Grimm, Escera, 2013, EJN

Early detection of violation within pattern regularities 14 A: Global deviant confirming local rule of frequency repetition breaking global

14 Early detection of violation within pattern regularities 14 A: Global deviant confirming local rule of frequency repetition breaking global pattern rule B: Local deviant breaking local rule of frequency repetition confirming global pattern rule A: No MLR effect! B: MLR effect! - Nb mmlr

breaking global pattern rule B: Local deviant breaking local rule of

15 Early detection of violation within pattern regularities 15 A: Global deviant confirming local rule of frequency repetition breaking global pattern rule B: Local deviant breaking local rule of frequency repetition confirming global pattern rule A: MMN! B: MMN/N1 effect mmmn

16 16 Whereas simple regularities are encoded in the earlier time range, more complex regularities (such as the frequent occurrence of specific patterns or feature combinations) are likely to be encoded only at the higher levels. (well, at least deviations of this simple and complex regularities are detected at different time points)

17 Can we observe deviance-related responses at the human scalp that reflect a processing of stimulus statistics at subcortical levels? 17

18 Deviance-related changes at subcortical levels? fmri 18 Stimuli were arranged in 2 different sessions Each session: 250 trials; each trial: 20 stimuli (constant SOA = 150ms) In total, 5000 stimuli delivered per session 12 subjects (age 24-36; 7 females; 4 left-handed) Session #1: 125 standard trials deviant trials Freq Freq Trial #1 Trial #2 Session #2: 125 control trials silent trials (baseline) Freq.5 Freq.4 Freq.3 Freq.2 Freq.1

19 Deviance-related changes at subcortical levels? fmri IC DEV>STA DEV>STD MGB IC DEV>CON MGB Cacciaglia et al., in preparation

Summary 20 deviance-related activity can be observed at

activities are organized in spatially distinct areas of the

indicate the early processing of simple feature changes, more

time range already at the level of the inferior colliculus, the

20 Summary 20 deviance-related activity can be observed at multiple time scales sources of these deviance-related activities are organized in spatially distinct areas of the Human Auditory Cortex whereas deviance-related MLR effects indicate the early processing of simple feature changes, more complex regularity violations are only reflected in the MMN time range already at the level of the inferior colliculus, the statistical status of a sound is processed as reflected by an enhanced BOLD response to deviants

21 Summary 21 Deviance detection, based on regularity encoding, is a key principle of the functional organization of the auditory system, spanning from lower levels in the auditory brainstem to higher order areas of the cerebral cortex. At higher levels in the auditory system deviant responses become stronger (in terms of amplitude) and functional complexity of deviant processing increases.

22 Thank you for your attention Thanks to the members of the Group for Cognitive Neuroscience at the University Barcelona: and to our collaborators: Kimmo Alho, Torsten Baldeweg, Manolo Malmierca, Israel Nelken, Minna Huotilainen, Erich Schröger, Istvan Winkler et al.

23 Important announcement: at the University of Leipzig, Germany

Detection of Simple and Pattern Regularity Violations Occurs at Different Levels of the Auditory Hierarchy

Detection of Simple and Pattern Regularity Violations Occurs at Different Levels of the Auditory Hierarchy Miriam Cornella 1,2, Sumie Leung 1,2, Sabine Grimm 1,2, Carles Escera 1,2 * 1 Institute for Brain,