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Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Max Planck Institute for Human Cognitive and Brain Sciences Leipzig, Germany Grounding language on neurobiology Angela D. Friederici
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Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

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Page 1: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Max Planck Institute for Human Cognitive and Brain Sciences Leipzig, Germany

Grounding language on neurobiology

Angela D. Friederici

Page 2: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Fitch.ppt

courtesy of T. Fitch

The language faculty is human specific. What is the neurobiological basis of

this faculty?

Page 3: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Friederici, Physiological Reviews, 2011

Fachbeirat_2012_NP.ppt

LH RH

Language in the human brain

Language is realized in the human brain as computations in specialized cortical areas that are tightly coupled functionally and structurally to form a large scale network for language processing. It receives its specificity due to syntax, a system of rules that permits the combination, organization and permutation of words in meta-structures, i.e. sentences.

Structure Structure Function Function

Page 4: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Pattern for non-embedded sentences

Pattern for embedded sentences

Basic neurophysiological principles

Amsterdam, 29.08.2014 >> Bilder >> Meyer.ppt

Although human specific, language is based on common neurophysiological principles. Individual neurons and neuronal assemblies are the basis of neuronal activity which for language usually can only be measured as spatiotemporal pattern at the cortical level (except for intracranial recordings).

Source: Meyer, Grigutsch, Schmuck, Gaston & Friederici, Poster presented at CNS, 2014

Page 5: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

160 ms

550 ms

700 ms

ELAN

N400

P600 -3.0

3.0 µV

-2.0

2.0 µV

-3.0

3.0 µV

access to syntactic category info & phrase structure building

access to lexical-semantic info &

thematic role assignment

Tim

e

integration

Source: 3-Phases Model by Friederici, Brain and Language, 1995; Trends in Cognitive Sciences, 2002

EEG:Neurochronometry of language comprehension

EEG data have allowed to identify specific ERP components reflecting different aspects and levels during on-line sentence comprehension.

Page 6: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Localization of the ELAN effect

left hemisphere right hemisphere

#1

#2

#3

#4

#5

158 ms

142 ms

133 ms

143 ms

139 ms

#3

133 ms left hemisphere right hemisphere

Source: Friederici et al., Human Brain Mapping, 2000

Phase 1: MEG localization of the ELAN

Using MEG the ELAN representing initial phrase structure processes could be localized in the anterior temporal and inferior frontal cortex.

Page 7: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Phase 2: The N400 allows to differentiate different aspects

Source: Hagoort, Hald, Bastiaansen & Petersson, Science, 2004

Semantic N400-effect World knowledge N400-effect

During sentence comprehension semantic and world knowledge are rapidly integrated in parallel.

correct: The Dutch trains are yellow and very crowded. world knowledge violation: The Dutch trains are white and very crowded. semantic violation: The Dutch trains are sour and very crowded.

(300–550 ms) (300–550 ms)

Page 8: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Gunter, Friederici & Schriefers, Journal of Cognitive Neuroscience, 2000

Amsterdam 29.08. 2014 >> Bilder >> Gunter

Phase 3: The integration phase: P600

Semantic and syntactic information interact during integration phase. The syntactic P600 is modulated by low cloze probability.

gender agreement gender disagreement

High cloze probability Low cloze probability

Page 9: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Binder et al., Cerebral Cortex, 2009

Amsterdam 29.08. 2014 >> Bilder >> Gunter

+46 –46

The semantic network

The semantic network has been demonstrated to be of considerable size, involving inferior frontal and temporal regions in both hemispheres.

Left hemisphere Right hemisphere

Page 10: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Friederici et al., Cerebral Cortex, 2006

3.09

left hemisphere

The syntactic network: Syntactic complexity

Functional subparts of the syntactic network are well localizable when well controlled stimulus material is used.

Broca's area Complexity-activation

Page 11: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences Source: Zaccarella & Friederici, Poster presented at FENS, 2013

The syntactic network: Merge

Merge, the most basic syntactic operation of binding two elements together to form a phrase can be localized in a subpart of Broca's area, the ventral-anterior region C3.

Mass localization with cluster overlap

PH>LS activation Volume-of-interest

Amsterdam, 29.08.2014 >> Bilder >>Zaccarella

Page 12: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Orange: main effect of hierarchy Light Blue: main effect of distance Red: interaction

2103_CNS_2009.ppt: 050309_neues Bild_100408 Fig 2_V3

Yellow: BA 45

Source: Makuuchi, Bahlmann, Anwander & Friederici, PNAS, 2009

Local syntactic network

Local syntactic network consist of BA 44 (syntactic hierarchy) and inferior frontal sulcus (syntactic WM). These regions interact during the processing of hierarchically structured sentences.

Processing embedded sentences

Green: BA

Page 13: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Functional activation Structural connectivity

2511_bahlmann_fig3.fh11 2505_Leipzig_2006.ppt, umgefärbt: TractographyData.jpg

Source: Friederici, Bahlmann et al., PNAS, 2006

from BA44 to STG BA 44

Source: Friederici et al., NeuroReport, 2009

Long-range syntactic network

Long-range syntactic network consists of BA 44 (syntactic hierarchy), and pSTG/STS (integration).

BA44 pSTG/STS

Page 14: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

29.08.2014 Amsterdam >>Bilder >> Meyer

Sources: Meyer, Grigutsch, Schmuck, Gaston & Friederici, Poster presented at CNS, 2014

The frontal and parietal/temporal region involved in syntactic processing synchronize (theta oscillations) during processing of syntactically complex sentences.

Neural synchronization within the syntactic network

Theta synchronization

Source localization Left frontal-parietal coherence

Page 15: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Syntax (LH) – Prosody (RH) – Interaction (CC)

Source: Friederici, Kotz & von Cramon, Neuron, 2007

Controls

effect

Anterior CC lesion

effect

Posterior CC lesion

pros correct pros incorrect

Patients with lesions in the posterior Corpus Callosum do not show mismatch effect indicating that prosodic information (RH) cannot misguide the syntactic parser (LH).

–6 µV

0.8 s

PZ

6

–6 µV

0.8 s

PZ

6

–6 µV

0.8 s

PZ

6

Page 16: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Sources: Zilles, Bacha-Trams, Palomero-Gallogher, Amunts & Friederici, Cerebral Cortex, 2015

29.08.2014 Amsterdam >>Bilder >> Bacha Trams

Areas that are part of the larger language network (red) may be neuroreceptorarchitectonically similar thereby providing a molecular basis for large-scale interaction.

The language network at the molecular level

Distribution of neuro-receptors Red: Language network

Page 17: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Hierarchical cluster tree Multidimensional scaling

Sources: Zilles, Bacha-Trams, Palomero-Gallogher, Amunts & Friederici, Cerebral Cortex, 2015

29.08.2014 Amsterdam >>Bilder >> Bacha Trams

Areas that are part of the language network show a more similar multi-receptor organisation compared to other regions.

The language network at the molecular level

Page 18: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Conclusion I

All the different levels of neurobiological analyses are relevant for our understanding of the brain-language relationship. Specific levels of description and their analytic methods should be applied for specific (linguistic) questions. But unless these level-specific analyses stay informed about the other levels, they will not advance science in the neurobiology of language.

Page 19: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Conclusion II

Unless we – as those being interested in the brain-language relationship – do not learn to consider more than one level of neurobiological description, the scientific community will not recognize the importance of our work, which in the end is crucial for our understanding of what it means to be human.

Page 20: Grounding Language on Neurobiology · Grounding language on neurobiology Angela D. Friederici . Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences

Amsterdam, August 29, 2014 Max Planck Institute for Human Cognitive and Brain Sciences