Analyse von 82 Hirnaktivierungsxperimenten mit vier verschiedenen Wortproduktionsaufgaben: Bildbenennung Wortgenerierung (z.B. Nennen Sie möglichst viele Tiere!) Wortlesen (HUND) Pseudowortlesen (HUNG)

Talairach & Tournoux (1988) Lateral and medial view of reference brain

Reported at least once Reported at least once

Estimate of probability of overlap under the assumption of a random distribution of activated regions number of regions: 110 mean number of activated regions: r chance probability for a region to be reported as activated in a single experiment (p1): r/110 chance probability for a region to be reported as activated in n1 out of n experiments: (with n1 + n2 = n)

Reliability criterion: p < 0 Reliability criterion: p < 0.1 cut-off point in binomial distribution Example region 1 Number of experiments: 82 Mean number of reported regions: 12.4 Reliably activated: 12 or more experiments Reliably not activated: 4 or less experiments Example region 2 Number of experiments: 23 Mean number of reported regions: 10.4 Reliably activated: 5 or more experiments Reliably not activated: -

Zuverlässig aktivierte (rot) und nicht aktivierte (blau) Hirngebiete (basierend auf allen 82 Studien)

TASK ANALYSIS Many tasks were not just word production tasks; they involved other operations as well. For instance, when you name the picture of a horse, you not only produce the word 'horse', but you also look at the picture and recognize it. Such additional 'lead-in' operations involve the activation of additional brain regions. These should be filtered out. That requires a systematic task analysis, a distinction between 'lead-in' and 'core' operations of word production.

Responses during Verb Generation Task BANANA TROUSERS CHAIR GLASSES TRUMPET PENCIL BUTTON BIRD EAR DOOR peel, slip on, eat up, plant put on, wash, mend, buy, warm sit, build, nail, sell, work, learn clean, put on, step on, buy, see blow, make music, put away, hear, play sharpen, break, put away, draw tear off, close, open fly, eat up, sing hear, pinch open, close, kick against

Aufgabe Einleitungsprozesse Kernprozesse Selbstmonitoring visuelle Objekt-erkennung Bildbenennung Konzeptuelle Vorbereitung lexikalische Selektion lexikalisches Konzept Lemma Wortformzugriff Wortform Syllabifizierung phonologisches Wort phonetische Enkodierung abstraktes Motorprogramm Artikulation gesprochenes Wort Selbstmonitoring Worterkennung Objektvorstellung Gedächtnis etc. Wortgenerierung visuelle Worterkennung Wortlesen Graphem/PhonemKonversion Pseudowortlesen aussprechen vs. Wort “denken”

Bildbenennung

Wortgenerierung

Bildbenennung (grün), Wortgenerierung (blau), gemeinsame Gebiete (rot)

Gemeinsame Aktivierungsgebiete von Bildbenennung und Wortgenerierung

Aufgabe Einleitungsprozesse Kernprozesse Selbstmonitoring visuelle Objekt-erkennung Bildbenennung Konzeptuelle Vorbereitung lexikalische Selektion lexikalisches Konzept Lemma Wortformzugriff Wortform Syllabifizierung phonologisches Wort phonetische Enkodierung abstraktes Motorprogramm Artikulation gesprochenes Wort Worterkennung Objektvorstellung Gedächtnis etc. Wortgenerierung visuelle Worterkennung Selbstmonitoring Wortlesen Graphem/PhonemKonversion Pseudowortlesen aussprechen vs. Wort “denken”

Aufgabe Einleitungsprozesse Kernprozesse Selbstmonitoring visuelle Objekt-erkennung Bildbenennung Konzeptuelle Vorbereitung lexikalische Selektion lexikalisches Konzept Lemma Wortformzugriff Wortform Syllabifizierung phonologisches Wort phonetische Enkodierung abstraktes Motorprogramm Artikulation gesprochenes Wort Worterkennung Objektvorstellung Gedächtnis etc. Wortgenerierung visuelle Worterkennung Selbstmonitoring Wortlesen Graphem/PhonemKonversion Pseudowortlesen aussprechen vs. Wort “denken”

Wortlesen

Gemeinsame Aktivierungsgebiete von Bildbenennung, Wortgenerierung und Wortlesen

Aufgabe Einleitungsprozesse Kernprozesse Selbstmonitoring visuelle Objekt-erkennung Bildbenennung Konzeptuelle Vorbereitung lexikalische Selektion lexikalisches Konzept Lemma Wortformzugriff Wortform Syllabifizierung phonologisches Wort phonetische Enkodierung abstraktes Motorprogramm Artikulation gesprochenes Wort Worterkennung Objektvorstellung Gedächtnis etc. Wortgenerierung visuelle Worterkennung Selbstmonitoring Wortlesen Graphem/PhonemKonversion Pseudowortlesen aussprechen vs. Wort “denken”

Gemeinsame Aktivierungsgebiete aller Aufgaben

Aussprechen im Vergleich zu Wort “denken”

Schematische Darstellung des Ergebnisses der Meta-Analyse von 82 Hirnaktivierungsstudien Indefrey, P. and Levelt, W.J.M. (2004) Cognition

The cognitive architecture of listening to language integration with other knowledge sources interpretation syntactic analysis thematic analysis word recognition phonological processing phonemes, syllables segmenting speech code decoding speech signal

Tekst Sereno Then once you have examined the city you can get a uh nice contrast to the surrounding country side - uh a very unique country side which contrasts the distinction between the the mountains to the uh low land of the coastal regions where there is a lot more uh fishing. Speech signal

snelheid proposities (rate of propositions) 1 2 3 4 5 6 7 8 9 10 seconds snelheid proposities (rate of propositions) snelheid lexical access (rate of words) snelheid klanken (rate of phonemes)

mixing van alle vier speech signal rate of propositions rate of words rate of phonemes

Reversed speech versus silence

Word lists versus silence

Studies comparing auditory stimuli to silent baseline conditions Study Stimulus # Belin 1998 200ms frequency transition, 60/min 1 40ms frequency transition, 60/min 2 Belin 1999 synthetic diphthong, 6/min 3 Binder 2000 tones, different frequencies, 90/min 4 Bookheimer 1998 pseudowords, 9/min 5 Celsis 1999 syllables, 180/min 6 tones, 500 + 700Hz, 180/min 7 di Salle 2001 tones, 1000Hz, 6/min 8 Engelien 1995 environmental sounds, 10/min 9 Fiez 1996 pseudowords, 60/min 10 words, 60/min 11 Giraud 2000 vowels vs. expecting vowels, 120/min 12 Holcomb 1998 tones, 1500Hz + lower tones, 30/min 13 Jäncke 1999 tones, 1000Hz, 60/min 14 Lockwood 1999 tones, 500 + 4000Hz, 60/min 15 Mellet 1996 words, 30/min 16 Mirz 1999 music 17 sentences 18 Study Stimulus # Mirz 1999 tones, 1000Hz 19 tones, 1000 + 4000Hz 20 words 21 Müller 1997 sentences, 12/min 22 Petersen 1988 words, 60/min 23 Price 1996 words, 40/min 24 words, different rates 25 Suzuki 2002a 26 Suzuki 2002b tones, 1000Hz, 60/min 27 Thivard 2000 tones with spectral maxima, 60/min 28 Warburton 1996 words, 4/min 29 Wise 1991 pseudowords, 40 or 60/min 30 Wong 1999 reversed sentences, 30/min 31 sentences, 30/min 32 words, 30/min 33 Wong 2002 reversed words, 15/min 34 35 words, 15/min 36 Indefrey & Cutler, 2004

Studies comparing auditory stimuli to simpler auditory stimuli Study Stimulus vs. control stimulus # Benson 2001 CVC > CV > V 1 Binder 1996 words vs. tones 2 Binder 2000 pseudo vs. tones 3 reversed words vs. tones 4 5 Giraud 2000 amplitude modulated noise vs. noise 6 sentences vs. vowels 7 words vs. vowels 8 Hall 2002 frequency modulated vs. static tone 9 harmonic vs. single tone 10 Jäncke 2002 syllables vs. 350 ms white noise bursts 11 syllables vs. steady state portion of vowel 12 syllables vs. tones 13 Müller 2002 90% 1000Hz + 10% 500Hz vs. 1000Hz 14 Mummery 1999 words vs. signal correlated noise 15 Price 1996 words vs. reversed words 16 Schlosser 1998 sentences vs. unknown language 17 Scott 2000 sentences vs. rotated sentences 18 Thivard 2000 frequency transition vs. stationary tone 19 Indefrey & Cutler, 2004

Talairach & Tournoux (1988) Lateral and medial view of reference brain

Silent control

Silent control

When do activation maxima agree reliably between studies?

Silent control

Silent control

Silent control

Narain et al. 2003, Fig. 2

Silent control

Silent control

Silent control

Silent control

Silent control

What about the functional roles of these areas?

Silent control

Auditory control

Auditory control

Silent control

Silent control

Summary Listening to speech without an additional task induces extensive bilateral temporal activation but no reliable activation of Broca’s area.

Summary With increasing linguistic complexity of stimuli, the distance of activation maxima from the primary auditory cortex increases; particularly in the left hemisphere. It seems to be the highest linguistic processing level that leads to the most significant activation difference compared to a silent control.

Summary The left hemisphere shows a clearer stimulus-specific differentiation of activation maxima. Areas that seem to be especially related to (post-) lexical and sentence level processing can be identified.

Summary bilateral posterior STG: phonology left posterior STS: lexical phonology left anterior STS: possibly lexical and sentential prosody, possibly lexical and sentential meaning

Hagoort & Indefrey, in press

Neuroimaging studies on sentence processing Hagoort & Indefrey, in press

Haller, Klarhöfer, Radue, Schwarzbach, & Indefrey (2007) Eur. J Haller, Klarhöfer, Radue, Schwarzbach, & Indefrey (2007) Eur. J. Neuroscience

Stimuli

Haller, Klarhöfer, Radue, Schwarzbach, & Indefrey (2007) Eur. J Haller, Klarhöfer, Radue, Schwarzbach, & Indefrey (2007) Eur. J. Neuroscience

Bookheimer (2002), Fig. 2

Haller, Klarhöfer, Radue, Schwarzbach, & Indefrey (2007) Eur. J Haller, Klarhöfer, Radue, Schwarzbach, & Indefrey (2007) Eur. J. Neuroscience

wegstossen-Animation(1)

wegstossen-Animation(2)

Der rote Kreis stößt die grüne Ellipse weg. Condition1: Sentences Der rote Kreis stößt die grüne Ellipse weg. (The red circle pushes the green ellipse away.) Condition 2: Noun phrases roter Kreis, grüne Ellipse, wegstoßen (red circle, green ellipse, push away) Condition 3: Single words Kreis, rot, Ellipse, grün, wegstoßen (circle, red, ellipse, green, push away) All conditions at slow (6/min) and fast (8/min) rate.

Sentences vs. Single Words Activation maximum at -54,6,10 Indefrey et al. (2001) PNAS Activation maximum at -60,14,12 Indefrey et al. (2004) Brain & Language

S and NP production vs. control (W) Indefrey, Hellwig, Herzog, Seitz & Hagoort (2004) Brain & Language

Conclusions (1) The left posterior IFG and the left posterior temporal lobe subserve syntactic comprehension. Neural activation in syntactic comprehension depends on the need for syntactic analysis. The two areas do not subserve the same function, because the temporal area does not seem to respond to syntactic errors and is not found in syntactic production.

Aufgabe vom 14.5.10 Finden Sie eine neue Studie (ab 2006) in der mit FMRI, PET, oder NIRS entweder Wortproduktion oder Wortverstehen oder Satzverstehen untersucht wurde. Vergleichen Sie die Ergebnisse mit der entsprechenden Meta-analyse. Wodurch könnten Unterschiede zustande gekommen sein?