Brain Imaging of Language Functions

Brain Imaging of Language Functions

Course number: 27-503-01

Lecturer: Prof. Michal Ben-Shachar

First Semester

Hours: 4 h/w – 2 credit points


Class requirements:

  • Attendance (over 80% attendance required)
  • Active attendance (read and discuss papers) AND class presentation (20% of final grade).
  • Final exam (exam material includes the contents of the lectures and final list of required reading; 80% of final grade).



Day 1

Day 2

Required reading



Cognitive neuropsychology, Modularity

  • Pinker, 1991
  • Small & Hickok, NBoL Ch1


Stroke Aphasia

Primary Progressive Aphasia

  • Dronkers et al., 2004
  • Henseler et al., 2014
  • Forster 1999



Lexical representation and lexical access

  • Mesulam et al., 2021
  • Baldo et al., 2018
  • Andrews et al., 2022
  • Thompson et al., 2015
  • Davis, NBoL Ch44*


Class presentations

Class presentations

  • Mesulam et al., 2015
  • Matchin et al., 2022
  • Pinker and Ullman, 2002
  • Leminen et al., 2018*



Class presentations

  • Bozic et al., 2013
  • Carota et al., 2016




  • Shapiro, 1997*
  • Sprouse & Hornstein, Ch14*


fMRI of morphological processing

Class presentations

  • Hauser, Chomsky & Fitch 2002*
  • Altmann, 1998


Online sentence processing

fMRI of syntactic processing

  • Matchin et al., 2017
  • Wilson et al., 2018


ERP of sentence processing

Class presentations


  • Hahne and Friederici, 1999
  • Cohen and Dehaene 2011
  • Price and Devlin 2011



fMRI of reading

  • Glezer et al., 2015
  • Schuster et al., 2021


Class presentations

Class presentations

  • Lerma-Usabiaga et al., 2018
  • Bouhali et al., 2019


Diffusion MRI

dMRI of reading pathways

  • Frey et al., 2008
  • Floel et al., 2009


Class presentations

Wrap up and overflow

  • Ben-Shachar et al, 2007
  • Hickok and Poeppel, Ch25




* - Recommended reading, beyond the scope


Papers for class presentation:

  • Andrews JP, Cahn N, Speidel BA, Chung JE, Levy DF, Wilson SM, Berger MS, Chang EF. Dissociation of Broca's area from Broca's aphasia in patients undergoing neurosurgical resections. J Neurosurg. 2022 Aug 5:1-11.
  • Baldo JV, Kacinik N, Ludy C, Paulraj S, Moncrief A, Piai V, Curran B, Turken A, Herron T, Dronkers NF. Voxel-based lesion analysis of brain regions underlying reading and writing. Neuropsychologia. 2018 Jul 1;115:51-59.
  • Bouhali F, Bézagu Z, Dehaene S, Cohen L. A mesial-to-lateral dissociation for orthographic processing in the visual cortex. Proc Natl Acad Sci U S A. 2019 Oct 22;116(43):21936-21946.
  • Bozic M, Tyler LK, Su L, Wingfield C, Marslen-Wilson WD. Neurobiological systems for lexical representation and analysis in English. J Cogn Neurosci. 2013 Oct;25(10):1678-91.
  • Carota F, Bozic M, Marslen-Wilson W. Decompositional Representation of Morphological Complexity: Multivariate fMRI Evidence from Italian. J Cogn Neurosci. 2016 Dec;28(12):1878-1896.
  • Flöel A, de Vries MH, Scholz J, Breitenstein C, Johansen-Berg H. White matter integrity in the vicinity of Broca's area predicts grammar learning success. Neuroimage. 2009 Oct 1;47(4):1974-81.
  • Frey S, Campbell JS, Pike GB, Petrides M. Dissociating the human language pathways with high angular resolution diffusion fiber tractography. J Neurosci. 2008 Nov 5;28(45):11435-44.
  • Glezer LS, Kim J, Rule J, Jiang X, Riesenhuber M. Adding words to the brain's visual dictionary: novel word learning selectively sharpens orthographic representations in the VWFA. J Neurosci. 2015 Mar 25;35(12):4965-72.
  • Lerma-Usabiaga G, Carreiras M, Paz-Alonso PM. Converging evidence for functional and structural segregation within the left ventral occipitotemporal cortex in reading. Proc Natl Acad Sci U S A. 2018 Oct 16;115(42):E9981-E9990. 
  • Matchin W, Hammerly C, Lau E. The role of the IFG and pSTS in syntactic prediction: Evidence from a parametric study of hierarchical structure in fMRI. Cortex. 2017 Mar;88:106-123.
  • Matchin W, den Ouden DB, Hickok G, Hillis AE, Bonilha L, Fridriksson J. The Wernicke conundrum revisited: evidence from connectome-based lesion-symptom mapping. Brain. 2022 Nov 21;145(11):3916-3930.
  • Mesulam MM, Thompson CK, Weintraub S, Rogalski EJ. The Wernicke conundrum and the anatomy of language comprehension in primary progressive aphasia. Brain. 2015 Aug;138(Pt 8):2423-37.
  • Mesulam MM, Coventry CA, Rader BM, Kuang A, Sridhar J, Martersteck A, Zhang H, Thompson CK, Weintraub S, Rogalski EJ. Modularity and granularity across the language network-A primary progressive aphasia perspective. Cortex. 2021 Aug;141:482-496.
  • Schuster S, Himmelstoss NA, Hutzler F, Richlan F, Kronbichler M, Hawelka S. Cloze enough? Hemodynamic effects of predictive processing during natural reading. Neuroimage. 2021 Mar;228:117687.
  • Thompson HE, Robson H, Lambon Ralph MA, Jefferies E. Varieties of semantic 'access' deficit in Wernicke's aphasia and semantic aphasia. Brain. 2015 Dec;138(Pt 12):3776-92.
  • Wilson SM, Bautista A, McCarron A. Convergence of spoken and written language processing in the superior temporal sulcus. Neuroimage. 2018 May 1;171:62-74.


Supporting papers and reviews (not for student presentations):

  • Neurobiology of language, Book, Editors: Hickok and Small, Chapters 1, 13, 14, 25, 29, 44*.
  • Pinker S. (1991). Rules of language. Science 253(5019):530-5.
  • *Hauser, M.D., Chomsky, N. & Fitch, W.T. (2002). Science 298, 1569–1579.
  • Dronkers NF, Wilkins DP, Van Valin RD Jr, Redfern BB, Jaeger JJ. (2004). Lesion analysis of the brain areas involved in language comprehension. Cognition 92(1-2):145-77.
  • *Henseler I, Regenbrecht F, Obrig H. Lesion correlates of patholinguistic profiles in chronic aphasia: comparisons of syndrome-, modality- and symptom-level assessment. Brain. 2014 Mar;137(Pt 3):918-30. * Beyond the scope
  • Forster 1999. The Microgenesis of Priming Effects in Lexical Access. B&L.
  • *Leminen A, Smolka E, Duñabeitia JA, Pliatsikas C. (2018). Morphological processing in the brain: The good (inflection), the bad (derivation) and the ugly (compounding). Cortex. pii: S0010-9452(18)30266-1.
  • Altmann 1998. Ambiguity in sentence processing. TICS.
  • Ben-Shachar, Dougherty and Wandell 2007. White matter pathways in reading. Current opinions in Neurobiology.
  • Hagoort 2008. The fractionation of spoken language understanding by measuring electrical and magnetic brain signals. Phil. Trans Royal Soc.
  • Shapiro 1997. Tutorial: an introduction to syntax.
  • Pinker and Ullman 2002. The past and future of the past tense. TICS.
  • Hahne and Friederici (1999). Electrophysiological Evidence for Two Steps in Syntactic Analysis: Early Automatic and late Controlled Processes. JoCN.
  • Dehaene S, Cohen L. The unique role of the visual word form area in reading. Trends Cogn Sci. 2011 Jun;15(6):254-62.
  • Price CJ, Devlin JT. The interactive account of ventral occipitotemporal contributions to reading. Trends Cogn Sci. 2011 Jun;15(6):246-53.


* Beyond the scope