<?xml version="1.0" encoding="ISO-8859-1"?><cms:container xmlns:cms="http://edoc.hu-berlin.de/diml/module/cms"><cms:document><cms:meta><cms:entry ref="front" type="front"/><cms:entry type="title">The neural correlates of human vibrotactile working memory: Converging evidence from functional magnetic resonance imaging, electroencephalographic and behavioral studies</cms:entry><cms:entry type="author">Claudia Preuschhof</cms:entry><cms:entry id="chapter1" part="chapter1" ref="chapter1" type="chapter">1</cms:entry><cms:entry id="N1008F" part="chapter1" ref="N1008F" type="section">1.1</cms:entry><cms:entry id="N10096" part="chapter1" ref="N10096" type="citenumber">1</cms:entry><cms:entry id="N100A9" part="chapter1" ref="N100A9" type="section">1.2</cms:entry><cms:entry id="N100AE" part="chapter1" ref="N100AE" type="subsection">1.2.1</cms:entry><cms:entry id="N100D4" part="chapter1" ref="N100D4" type="subsection">1.2.2</cms:entry><cms:entry id="N100DB" part="chapter1" ref="N100DB" type="citenumber">2</cms:entry><cms:entry id="N1013B" part="chapter1" ref="N1013B" type="citenumber">3</cms:entry><cms:entry id="N1019B" part="chapter1" ref="N1019B" type="citenumber">4</cms:entry><cms:entry id="N101A6" part="chapter1" ref="N101A6" type="section">1.3</cms:entry><cms:entry id="N101C8" part="chapter1" ref="N101C8" type="citenumber">5</cms:entry><cms:entry id="N101CE" part="chapter1" ref="N101CE" type="mm">485#302</cms:entry><cms:entry id="N101E0" part="chapter1" ref="N101E0" type="citenumber">6</cms:entry><cms:entry id="N1024A" part="chapter1" ref="N1024A" type="section">1.4</cms:entry><cms:entry id="N1024F" part="chapter1" ref="N1024F" type="subsection">1.4.1</cms:entry><cms:entry id="N10282" part="chapter1" ref="N10282" type="citenumber">7</cms:entry><cms:entry id="N102AB" part="chapter1" ref="N102AB" type="citenumber">8</cms:entry><cms:entry id="N102BD" part="chapter1" ref="N102BD" type="subsection">1.4.2</cms:entry><cms:entry id="N102CB" part="chapter1" ref="N102CB" type="citenumber">9</cms:entry><cms:entry id="N10310" part="chapter1" ref="N10310" type="subsection">1.4.3</cms:entry><cms:entry id="N10317" part="chapter1" ref="N10317" type="citenumber">10</cms:entry><cms:entry id="chapter2" part="chapter2" ref="chapter2" type="chapter">2</cms:entry><cms:entry id="N10365" part="chapter2" ref="N10365" type="helpercitenumber">10</cms:entry><cms:entry id="N10370" part="chapter2" ref="N10370" type="citenumber">11</cms:entry><cms:entry id="N1037F" part="chapter2" ref="N1037F" type="citenumber">12</cms:entry><cms:entry id="chapter3" part="chapter3" ref="chapter3" type="chapter">3</cms:entry><cms:entry id="N10391" part="chapter3" ref="N10391" type="helpercitenumber">12</cms:entry><cms:entry id="N10394" part="chapter3" ref="N10394" type="section">3.1</cms:entry><cms:entry id="N1039B" part="chapter3" ref="N1039B" type="citenumber">13</cms:entry><cms:entry id="N103B2" part="chapter3" ref="N103B2" type="mm">378#78</cms:entry><cms:entry id="N103BE" part="chapter3" ref="N103BE" type="section">3.2</cms:entry><cms:entry id="N103E4" part="chapter3" ref="N103E4" type="citenumber">14</cms:entry><cms:entry id="N103FE" part="chapter3" ref="N103FE" type="section">3.3</cms:entry><cms:entry id="N10449" part="chapter3" ref="N10449" type="section">3.4</cms:entry><cms:entry id="chapter4" part="chapter4" ref="chapter4" type="chapter">4</cms:entry><cms:entry id="N10484" part="chapter4" ref="N10484" type="citenumber">15</cms:entry><cms:entry id="N10488" part="chapter4" ref="N10488" type="section">4.1</cms:entry><cms:entry id="N1048D" part="chapter4" ref="N1048D" type="subsection">4.1.1</cms:entry><cms:entry id="N104AA" part="chapter4" ref="N104AA" type="citenumber">16</cms:entry><cms:entry id="N104E5" part="chapter4" ref="N104E5" type="subsection">4.1.2</cms:entry><cms:entry id="N104EC" part="chapter4" ref="N104EC" type="citenumber">17</cms:entry><cms:entry id="N10518" part="chapter4" ref="N10518" type="citenumber">18</cms:entry><cms:entry id="N10529" part="chapter4" ref="N10529" type="subsection">4.1.3</cms:entry><cms:entry id="N10530" part="chapter4" ref="N10530" type="citenumber">19</cms:entry><cms:entry id="N1056A" part="chapter4" ref="N1056A" type="section">4.2</cms:entry><cms:entry id="N1056F" part="chapter4" ref="N1056F" type="subsection">4.2.1</cms:entry><cms:entry id="N10576" part="chapter4" ref="N10576" type="citenumber">20</cms:entry><cms:entry id="N1059C" part="chapter4" ref="N1059C" type="subsection">4.2.2</cms:entry><cms:entry id="N105AA" part="chapter4" ref="N105AA" type="citenumber">21</cms:entry><cms:entry id="chapter5" part="chapter5" ref="chapter5" type="chapter">5</cms:entry><cms:entry id="N105CF" part="chapter5" ref="N105CF" type="helpercitenumber">21</cms:entry><cms:entry id="N105D2" part="chapter5" ref="N105D2" type="section">5.1</cms:entry><cms:entry id="N105D9" part="chapter5" ref="N105D9" type="citenumber">22</cms:entry><cms:entry id="N105FD" part="chapter5" ref="N105FD" type="section">5.2</cms:entry><cms:entry id="N1060D" part="chapter5" ref="N1060D" type="citenumber">23</cms:entry><cms:entry id="N1061D" part="chapter5" ref="N1061D" type="section">5.3</cms:entry><cms:entry id="N10652" part="chapter5" ref="N10652" type="section">5.4</cms:entry><cms:entry id="N10659" part="chapter5" ref="N10659" type="citenumber">24</cms:entry><cms:entry id="N10683" part="chapter5" ref="N10683" type="citenumber">25</cms:entry><cms:entry id="N1068F" part="chapter5" ref="N1068F" type="mm">563#677</cms:entry><cms:entry id="N1069A" part="chapter5" ref="N1069A" type="citenumber">26</cms:entry><cms:entry id="chapter6" part="chapter6" ref="chapter6" type="chapter">6</cms:entry><cms:entry id="N106A7" part="chapter6" ref="N106A7" type="helpercitenumber">26</cms:entry><cms:entry id="N106CA" part="chapter6" ref="N106CA" type="citenumber">27</cms:entry><cms:entry ref="N106D0" type="back"/><cms:entry id="N106D2" part="N106D2" ref="N106D2" type="acknowledgement">Danksagung</cms:entry><cms:entry id="N106ED" part="N106ED" ref="N106ED" type="declaration">Eidesstattliche Erklärung</cms:entry><cms:entry id="N106FE" part="N106FE" ref="N106FE" type="appendix">List of original publications</cms:entry><cms:entry id="N10700" part="N106FE" ref="N10700" type="head"/><cms:entry id="N10703" part="N106FE" ref="N10703" type="p"/><cms:entry id="N10706" part="N106FE" ref="N10706" type="p"/><cms:entry id="N1070F" part="N106FE" ref="N1070F" type="p"/><cms:entry id="N10712" part="N106FE" ref="N10712" type="p"/><cms:entry id="N1071B" part="N106FE" ref="N1071B" type="p"/><cms:entry id="N1071E" part="N106FE" ref="N1071E" type="p"/><cms:entry id="N10727" part="N106FE" ref="N10727" type="p"/><cms:entry id="N1072A" part="N106FE" ref="N1072A" type="p"/><cms:entry id="N10733" part="N106FE" ref="N10733" type="p"/><cms:entry id="N10736" part="N106FE" ref="N10736" type="p"/><cms:entry id="N1073F" part="N106FE" ref="N1073F" type="p"/><cms:entry 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type=":help"><url href="http://...">Hilfe</url></cms:entry></cms:meta><cms:content><front id="front"><title>The neural correlates of human vibrotactile working memory: Converging evidence from functional magnetic resonance imaging, electroencephalographic and behavioral studies</title><submission>Dissertation</submission><degree>zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.)<br/>im Fach Psychologie</degree><p>eingereicht an der</p><major/><major>Mathematisch-Naturwissenschaftlichen Fakultät II<br/>der Humboldt-Universität zu Berlin</major><p>vorgelegt von</p><author>Dipl. Psych. <given>Claudia</given> <surname>Preuschhof</surname><br/><suffix>geboren am 4. Juli 1975 in Berlin</suffix>
      </author><p>Präsident der Humboldt-Universität zu Berlin:<br/>Prof. Dr. Christoph Markschies</p><dean>Dekan der Mathematisch-Naturwissenschaftlichen Fakultät II:<br/>Prof. Dr. Wolfgang Coy</dean><approvals>
         <name>Prof. Dr. Elke van der Meer</name>
         <name>Prof. Dr. Torsten Schubert</name>
         <name>Dr. Hauke R. Heekeren</name>
         
      </approvals><date>Einreichungsdatum: 31.03.2008</date><date>Tag der Verteidigung: 28.11.2008</date><abstract lang="de">
         <head>Zusammenfassung</head>
         <p>Das Ziel der vorliegenden Arbeit war es, die Mechanismen und die neuronale Implementierung des taktilen Arbeitsgedächtnisses (AG) zu erleuchten. Dazu wurde eine vibrotaktile Diskriminationsaufgabe mit den Methoden der funktionellen Magnetresonanztomografie (fMRT), Elektroenzephalografie (EEG), subliminalen elektrischen Stimulation und Psychophysik untersucht. Eine fMRT-Studie (Studie I) zeigte, dass ein ausgedehntes neuronales Netzwerk mit den einzelnen Aufgabenphasen, Enkodierung, Aufrechterhaltung, und Entscheidung, verbunden ist. Die Analyse oszillatorischer EEG-Aktivität (Studie II) und die selektive Inhibition des primären somatosensorischen Kortex (S1) durch subliminale Stimulation (Studie III) legen nahe, dass S1 nicht an der Aufrechterhaltung der vibrotaktilen Gedächtnisrepräsentation beteiligt ist. Das Ausmaß der S1-Aktivierung während der frühen Haltephase hängt vielmehr von der Effizienz, mit welcher Versuchspersonen den vibrotaktilen Reiz enkodieren, ab. Außerdem weisen Studie II and III darauf hin, dass bereits der physiologische Zustand von S1 in der Phase vor Beginn eines Versuchsdurchgangs eine einflussreiche Rolle spielt. Während Studie II zeigte, dass S1-Aktivität in dieser Phase, vermutlich durch attentionale top-down Modulation, unter AG-Anforderungen erhöht ist, ergab Studie III, dass eine optimierte bottom-up Verarbeitung in S1 zu einer verbesserten Aufgabenleistung führt. Verhaltensexperimente zeigten, dass Repräsentationen im Langzeitgedächtnis über die durchschnittliche Frequenz der präsentierten Vibrationsreize erheblich die Verhaltensleistung beeinflussen und den Zeitrichtungseffekt verursachen (Studie IV). Die Analyse der fMRT-Daten (Studie V) offenbarte außerdem, dass ein Netzwerk somatosensorischer Areale Informationen über die aktuelle Vibrationsfrequenz und die durchschnittliche Vibrationsfrequenz während der Enkodier- und Haltephase integriert. </p>
      </abstract><keywords lang="de">
         <keyword>Schlagwörter: </keyword>
         <keyword>Somatosensorik</keyword>
         <keyword>vibrotaktil</keyword>
         <keyword>Arbeitsgedächtnis</keyword>
         <keyword>fMRT</keyword>
         <keyword>EEG</keyword>
      </keywords><abstract lang="en">
         <head>Abstract</head>
         <p>The present dissertation aimed to shed more light on the psychological mechanisms and the neural basis of tactile working memory (WM). For this purpose, a vibrotactile delayed discrimination task was studied using the methods of functional magnetic resonance imaging (fMRI), electroencephalography (EEG), concurrent subliminal electrical stimulation and psychophysics. The fMRI study (Study I) showed that a broad network of brain regions - much broader than known from previous studies in non-human primates - supports the performance of a vibrotactile delayed discrimination task in the different task periods: encoding, maintenance, decision making. The analysis of oscillatory activity over the somatosensory cortex in the EEG study (Study II) and the experiment using subliminal electrical stimulation to locally inhibit the primary somatosensory cortex (S1) (Study III) suggest that S1 does not contribute to the active maintenance of the vibrotactile memory trace. The level of activity in S1 during the early delay period depends on the efficiency with which subjects encode the vibrotactile stimulus. Study II and III also showed that the activation level of S1 in the pre-trial period plays an important role. Study III suggests that, in this task period, S1 activity is up-regulated under WM demands probably reflecting the operation of top-down attentional control. Study III indicates that increasing local inhibition of S1 in the pre-trial period improves performance by facilitating bottom-up processing. Importantly, long-term memory representations of the average frequency of the stimulus set strongly influence performance giving rise to the time-order effect (Study IV). Additionally, the fMRI data (Study V) showed that a somatosensory network integrates information about the current vibrotactile stimulus and the representation of the average vibration frequency during stimulus encoding and maintenance. </p>
      </abstract><keywords lang="en">
         <keyword>Keywords: </keyword>
         <keyword>somatosensory</keyword>
         <keyword>vibrotactile</keyword>
         <keyword>working memory</keyword>
         <keyword>fMRI</keyword>
         <keyword>EEG</keyword>
      </keywords><freehead id=":contents">Inhaltsverzeichnis</freehead><ul><li><p><link ref="chapter1">1</link> Theoretical background<ul><li><p><link ref="N1008F">1.1</link> Introduction</p></li><li><p><link ref="N100A9">1.2</link> The concept of working memory<ul><li><p><link ref="N100AE">1.2.1</link> Psychological concepts of working memory</p></li><li><p><link ref="N100D4">1.2.2</link> Neural basis of WM </p></li></ul></p></li><li><p><link ref="N101A6">1.3</link> The neuroanatomy of the somatosensory system</p></li><li><p><link ref="N1024A">1.4</link> Previous research on vibrotactile working memory<ul><li><p><link ref="N1024F">1.4.1</link> Single-unit studies of vibrotactile working memory in non-human primates </p></li><li><p><link ref="N102BD">1.4.2</link> Human studies on vibrotactile working memory</p></li><li><p><link ref="N10310">1.4.3</link> Time-order effects in vibrotactile working memory</p></li></ul></p></li></ul></p></li><li><p><link ref="chapter2">2</link> Summary of research questions</p></li><li><p><link ref="chapter3">3</link> Methodological background<ul><li><p><link ref="N10394">3.1</link> Vibrotactile delayed discrimination task</p></li><li><p><link ref="N103BE">3.2</link> Functional magnetic resonance imaging</p></li><li><p><link ref="N103FE">3.3</link> Electroencephalography</p></li><li><p><link ref="N10449">3.4</link> Subliminal electrical stimulation</p></li></ul></p></li><li><p><link ref="chapter4">4</link> Empirical studies <ul><li><p><link ref="N10488">4.1</link> Neural correlates of human vibrotactile working<ul><li><p><link ref="N1048D">4.1.1</link> Study I: Neural correlates of vibrotactile working memory in the human brain</p></li><li><p><link ref="N104E5">4.1.2</link> Study II: Rolandic and posterior-parietal alpha rhythms are differentially modulated during vibrotactile working memory </p></li><li><p><link ref="N10529">4.1.3</link> Study III: Pre-trial subliminal electrical stimulation improves performance in vibrotactile delayed discrimination </p></li></ul></p></li><li><p><link ref="N1056A">4.2</link> Time order effects in vibrotactile working memory: the influence of long-term memory representations on task performance<ul><li><p><link ref="N1056F">4.2.1</link> Study IV: Time-order effect in vibrotactile delayed discrimination: The influence of trial types and response alternatives</p></li><li><p><link ref="N1059C">4.2.2</link> Study V: Neural correlates of time order effects: Implicit sensory stimulus representations influence vibrotactile delayed discrimination</p></li></ul></p></li></ul></p></li><li><p><link ref="chapter5">5</link> General discussion<ul><li><p><link ref="N105D2">5.1</link> The neural network supporting vibrotactile WM</p></li><li><p><link ref="N105FD">5.2</link> The role of S1 in vibrotactile WM</p></li><li><p><link ref="N1061D">5.3</link>  The role of implicit LTM representations in vibrotactile WM</p></li><li><p><link ref="N10652">5.4</link> An integrated model for vibrotactile working memory</p></li></ul></p></li><li><p><link ref="chapter6">6</link> Conclusion</p></li><li><p><link ref="N106D2">Danksagung</link></p></li><li><p><link ref="N106ED">Eidesstattliche Erklärung</link></p></li><li><p><link ref="N106FE">List of original publications</link></p></li><li><p><link ref="N10743">List of Abbreviations</link></p></li><li><p><link ref="N108E1">Literature</link></p></li></ul><freehead id=":toc-media">Bilder</freehead><ul><li><p><link ref="N101CE">Figure 1. The somatosensory system. a) The somatosensory pathway from the mechanoreceptors to the primary (S1) and secondary (S2) somatosensory cortex. b) S1 comprises the postcentral gyrus in the anterior parietal lobe and contains the Brodman areas (BA) 3b, 3a, 1, 2 (see enlarged cutout). S1 projects to the ipsi- and contralateral S2, the primary motor cortex (M1), the inferior (IPL) and superior posterior parietal lobe (SPL). S2 is located in the upper bank of the Sylvian fissure and projects to contralateral S2, IPL, SPL, the ventral (vPMC) and medial premotor cortex (mPMC) and lateral prefrontal cortex (PFC).</link></p></li><li><p><link ref="N103B2">Figure 2. The vibrotactile delayed discrimination task. After a warning tone, two vibrotactile stimuli are presented successively to the subject's index finger. The subject has to discriminate between the two vibration frequencies.</link></p></li><li><p><link ref="N1068F">Figure 3. An integrated model of vibrotactile WM. a) summarizes the major claims of the different studies of this dissertation. b) shows the main regions of the neural network related to the different task periods. Please note that neural network supporting vibrotactile WM is much broader than depicted here. Abbreviations: S1, primary somatosensory cortex; S2, secondary somatosensory cortex; M1, motor cortex; mPMC, medial premotor cortex; vPMC, ventral premotor cortex; PFC, prefrontal cortex; IPL, inferior parietal lobe; SPL, superior parietal lobe. </link></p></li></ul></front><img src="http://vg01.met.vgwort.de/na/b7d4da7d3250989336a4c7c68b94ea" width="1" height="1" alt=""/></cms:content></cms:document></cms:container>