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Auditory Selective Attention: an Introduction and Evidence for Distinct Facilitation and Inhibition Mechanisms

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Auditory Selective Attention: an Introduction and Evidence for Distinct Facilitation and Inhibition Mechanisms Aus dem Epilepsiezentrum Erlangen Leiter: Prof. Dr. med. Hermann Stefan der Neurologischen Klinik mit Poliklinik der Friedrich-Alexander-Universität Erlangen-Nürnberg Direktor: Prof. Dr. med. Stefan Schwab Durchgeführt im Helen Wills Neuroscience Institute der University of California, Berkeley, USA Direktor: Robert T. Knight, M.D. Betreuer: Aurélie Bidet-Caulet, Ph.D. Auditory Selective Attention: an introduction and evidence for distinct facilitation and inhibition mechanisms Inaugural-Dissertation zur Erlangung der Doktorwürde der Medizinischen Fakultät der Friedrich-Alexander-Universität Erlangen-Nürnberg vorgelegt von Constanze Elisabeth Anna Mikyska aus München Gedruckt mit Erlaubnis der Medizinischen Fakultät der Friedrich-Alexander-Universität Erlangen-Nürnberg Dekan: Prof. Dr. med. Dr. h.c. J. Schüttler Referent: Prof. Dr. med. H. Stefan Korreferent: Prof. Dr. med. Dipl.-Psych. Ch. Lang Tag der mündlichen Prüfung: 29. Februar 2012 To my family Table of contents 1 Summary 1 1.1 Summary 1 1.2 Zusammenfassung 3 2 Introduction 5 2.1 Auditory system: anatomy and function 5 2.1.1 Ear 5 2.1.2 Sub-cortical auditory relays 10 2.1.3 Auditory cortex 13 2.2 Investigation of auditory perception and processing 14 2.2.1 Psychophysics – psychoacoustics 15 2.2.2 Brain activity – electroencephalography (EEG) 17 2.2.2.1 Introduction and history 17 2.2.2.2 Physiological fundamentals 17 2.2.2.3 Recording 22 2.2.2.4 Classification of frequency 25 2.2.2.5 Artifacts 27 2.2.2.6 Data analysis: preprocessing and event-related potentials (ERP) 30 2.2.2.7 Main auditory electrophysiological components 31 2.3 Auditory attention 35 2.3.1 Psychological theories 35 2.3.1.1 Introduction to selective attention 35 2.3.1.2 Bottleneck theories: early- versus late-selection 37 2.3.1.3 Other capacity-limitation theories 40 2.3.2 Electrophysiological findings and theories 42 2.4 Aims of this dissertation 49 3 Material and methods 54 3.1 Subjects 54 3.2 Stimuli and task 54 3.3 Procedure 57 3.4 EEG recording 57 3.5 EEG data analysis 57 3.6 Statistical analysis 58 3.6.1 Selection of applied methods 58 3.6.1.1 Analysis of variance (ANOVA) 58 3.6.1.2 Statistic permutation test 59 3.6.2 Behavioral data 60 3.6.3 ERP standards 61 3.6.4 ERP deviants 62 4 Results 64 4.1 Behavioral data 64 4.2 ERP results of standards 65 4.2.1 Main attention effect (attended versus ignored) 65 4.2.2 Influence of the memory task difficulty on attention effects 66 4.2.3 Timing of attention facilitation and inhibition 70 4.2.4 Topographies of attention facilitation and inhibition 72 4.3 ERP results of deviants 73 4.3.1 Attention enhancement of deviant processing 73 4.3.2 Memory effect on the P3-Component 75 5 Discussion 77 6 References 82 7 List of abbreviations 90 8 Publication 92 9 Acknowledgements 93 10 Curriculum vitae 94 1 1 Summary 1.1 Summary Objective Auditory selective attention is a complex brain function that is still not completely understood. The classic example is the so-called “cocktail party effect” (Cherry, 1953), which describes the impressive ability to focus one’s attention on a single voice from a multitude of voices. This means that particular stimuli in the environment are enhanced in contrast to other ones of lower priority that are ignored. To be able to understand how attention can influence the perception and processing of sound, background knowledge is essential. One aim of this dissertation is to provide an overview of already existing literature. Therefore, the auditory system and different methods to measure and evaluate auditory processes are introduced at first, followed by a review about competing theories, trying to explain how auditory attention operates. The second aim of the dissertation is to specify the mechanisms and to elucidate how they operate. It is generally accepted that distinct signals (top-down signals) are important for cognitive control, enabling selective attention and leading to an enhanced processing of task relevant information. But it is unknown whether facilitation and inhibition of stimulus processing are based upon one (unitary gain control mechanism of facilitation) or two mechanisms (net activity of distinct top- down facilitation and inhibition mechanisms). Results from a visual fMRI study (de Fockert, 2001) suggest that facilitation and inhibition rely on distinct mechanisms that would be differentially affected by the availability of cognitive resources (i.e. for performing a task). To reveal that facilitation and inhibition represent distinct mechanisms in auditory selective attention, we conducted a study, where subjects performed an auditory attention task, while the amount of available cognitive resources was modulated (by varying the difficulty of a memory task). Methods Electrophysiological experiments were conducted in young healthy adults. 16 subjects performed an attention task and a memory task of varying difficulty (no, easy and difficult memory) at the same time (dual task protocol) while EEG was recorded. Facilitation and inhibition were measured by comparing 2 electrophysiological responses to attended and ignored sounds with responses to the same sounds when attention was considered to be equally distributed towards all sounds. Results Two ERP-components were observed: a negative one in response to attended sound and a positive one to ignored sounds. The two frontally distributed components had distinct timing and scalp topographies and were differentially affected by the difficulty of the memory load. Conclusion This dissertation provides an insight into the literature of auditory selective attention and also enriches the existing knowledge with results of a new study about the operating mechanisms of auditory selective attention. The study provides evidence that top-down attention control can operate via distinct facilitation and inhibition mechanisms. 3 1.2 Zusammenfassung Hintergrund und Ziele Selektive auditorische Aufmerksamkeit ist ein komplexer Mechanismus, der noch nicht vollständig verstanden ist. Das klassische Beispiel ist der sogenannte „Cocktail Party Effekt“ (Cherry, 1953). Dieser beschreibt die beeindruckende Fähigkeit, die Aufmerksamkeit auf einen einzelnen Sprecher zu konzentrieren und andere Unterhaltungen auszublenden. Das bedeutet, dass bestimmte Reize in unserer Umwelt verstärkt wahrgenommen werden, wohingegen Reize von niedrigerer Priorität ignoriert werden. Um zu verstehen, wie Aufmerksamkeit die Wahrnehmung und Verarbeitung von Reizen beeinflusst, gibt der erste Teil dieser Dissertation einen Überblick der Grundlagenliteratur. Dabei werden zuerst das auditorische System vorgestellt und verschiedene Methoden zur Messung und Beurteilung auditorischer Verarbeitungsprozesse eingeführt. Dem folgt ein kurzer Überblick über konkurrierende Theorien, die zu erklären versuchen, wie selektive auditorische Aufmerksamkeit funktioniert. Der zweite Teil dieser Arbeit befasst sich genauer mit der Frage nach den Mechanismen und wie diese arbeiten. Es ist allgemein anerkannt, dass bestimmte Signale (top-down Signale) wichtig für die kognitive Kontrolle sind. Sie aktivieren selektive auditorische Aufmerksamkeit und führen so zu einer verstärkten Verarbeitung eines relevanten Reizes. Aber es ist noch ungeklärt ob die Förderung und Hemmung der Reizverarbeitung durch einen (einheitlicher, linearer Verstärkungsmechanismus von Förderung) oder zwei Mechanismen (Netzwerkaktivität von unabhängiger Förderung und Hemmung) geregelt wird. Ergebnisse einer visuellen fMRT Studie zeigen, dass das Ausmaß der Hemmung ablenkender Reize von der Verfügbarkeit kognitiver Ressourcen (z.B. für das Lösen von Problemen) abhängig ist (de Fockert, 2001). Die Ergebnisse deuten darauf hin, dass Förderung und Hemmung im visuellen System auf verschiedenen Mechanismen basieren, die von der Verfügbarkeit kognitiver Ressourcen unterschiedlich beeinflusst werden. Um zu zeigen, dass Förderung und Hemmung unabhängig voneinander agieren, führten wir eine Studie durch, in der Probanden einen auditorischen Aufmerksamkeitstest lösten, während die Verfügbarkeit von kognitiven Ressourcen variiert wurde (verschiedene Schwierigkeitsstufen in einem Gedächtnis Test). 4 Methoden Elektrophysiologische Versuche wurden mit 16 jungen, gesunden Erwachsenen durchgeführt. Die Probanden lösten gleichzeitig (dual task protocol) einen Aufmerksamkeits- und einen Gedächtnis Test mit variierenden Schwierigkeitsstufen (no, easy und difficult memory) während elektrophysiologische Signale (EEG) aufgezeichnet wurden. Förderung und Hemmung wurden gemessen, indem die Antworten zu den beachteten und den ignorierten Reizen jeweils mit den Antworten auf die gleichen Reize einer Kontrollbedingung verglichen wurden. In dieser Kontrollbedingung wurde angenommen, dass die Aufmerksamkeit ausgewogen auf alle Reize gerichtet war. Ergebnisse und Beobachtungen Zwei ERP-Komponenten wurden beobachtet: eine negative, in Antwort zu den beachteten Reizen und eine positive, den ignorierten Reizen folgend. Die zwei Komponenten zeigten verschiedene frontale Skalp-Topographien und variierten auch in der zeitlichen Domäne. Außerdem wurden sie unterschiedlich von der Schwierigkeit des Gedächtnis Tests beeinflusst. Praktische Schlussfolgerungen Diese Dissertation bietet einen Einblick in die Literatur über selektive auditorische Aufmerksamkeit und bereichert das bestehende Wissen mit Ergebnissen einer neuen Studie über die Wirkmechanismen. Die Studie erbringt den Nachweis, dass top-down Kontrolle die Aktivität voneinander unabhängiger Förderungs- und Hemmungsmechanismen widerspiegelt.
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