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The Neuroscience of
     Language
From Structure to Process
Neuroscience and Language
• Aphasia (or dysphasia) – Language
  disorder of auditory or oral speech,
  writing (agraphia), or reading (alexia)
  produced by injury to brain areas
  specialized for these functions
Nature’s Experiments
Nature’s Experiments
          • Stroke
            – Status post 5 hours
Nature’s Experiments
• Status post 5 days.
Blood-Brain Barrier
          • Blood is toxic to
            neurons.
             – Makes evolutionary
               sense.
             – Insulates brain from
               what’s in the blood.
Paul Broca
                 • Broca (1865)
                   described patients
                   who displayed
                   halting, agrammatic
                   speech
                   – Content words were
                     well preserved
Paul Broca
1824-1880          – Function words (i.e.,
                     adjectives, articles)
                     impaired
Broca’s Aphasia




• Patient “Tan”
• Brain tumor in Left frontal brain region
• Broca: Lesion disrupted speech
Broca’s Aphasia




• Broca’s Aphasia – Damage to “motor images”
• Language comprehension skills relatively
  preserved
• Typically observed in patients with damage to
  left inferior prefrontal cortex
Broca’s Aphasia
• “Yes… ah… Monday… er… Dad and Peter
  H… (patient’s name), and Dad… er…
  hospital… and ah… Wednesday…
  Wednesday, nine o’clock… and oh…
  Thursday… ten o’clock, ah doctors… two…
  an’ doctors… and er… teeth…yah

                     Goodglass & Geschwind, 1976
Carl Wernicke
      • Wernicke (1874)
        described patients
        whose speech is
        fluent, but has no
        informational value
Wernicke’s Aphasia
• Neologisms
• Speech appears to have no
  information content
• “fluent nonsense”
• Preserved function words,
  impaired content words
• Comprehension impaired
• Even simple sentences not well
  understood
• Associated with left temporal
  lobe damage
Wernicke’s Aphasia
• “Well this is… mother is away here working
  her work out o’here to get her better, but
  when she’s looking in the other part. One
  their small tile into her time here. She’s
  working another time…”



                    Goodglass & Geschwind, 1976
Articulatory        Auditory speech
speech “memories”      “memories”




                      Geschwind 1979
                     Scientific American
Articulatory        Auditory speech
speech “memories”      “memories”




 Production          Comprehension
 problems                   problems
                                +
                    empty error-filled
                          production
Broca’s aphasia   Wernicke’s aphasia

  Production        Comprehension
  problems                 problems
                               +
                   empty error-filled
                         production
Lichtheim
• Lichtheim (1885) – cases of patients
  able to understand and produce speech
  but unable to repeat words
• “Conduction aphasia”
• Hypothesis: Broca’s area (speech
  production) and Wernicke’s area
  (speech comprehension) intact
Conduction aphasia
      Error-filled speech, but
       good comprehension




Broca’s aphasia                  Wernicke’s aphasia

  Production                       Comprehension
  problems                                problems
                                              +
                                  empty error-filled
                                        production
Broca’s, Wernicke’s Area, and
               Connections




• Lichtheim’s (1885) and Geschwind’s (1965)
  model
• Auditory input mediated by Wernicke’s area
• Motor output mediated byBroca’s area
• Regions connected by arcuate fasciculus
Wernicke-Lichtheim “House” Model

                                Concept representations
                         C
                                  Widely distributed!




                    M               A
Motor word images                    Auditory word images
  Broca’s area                         Wernicke’s area
Lichtheim/Geschwind Model
                                Association Cortex

                          Concepts

Ventral prefrontal                         Posterior Temporal
cortex Motor word                     Auditory word Cortex
        Comprehension    Arcuate      Comprehension
                         Fasciculus


   Speech motor output
                                              Auditory input
Lichtheim/Geschwind Model

                          Concepts    Association Cortex




         X
Ventral prefrontal                        Posterior Temporal
cortex Motor word                     Auditory word Cortex
        Comprehension    Arcuate      Comprehension
                         Fasciculus


   Speech motor output
                                              Auditory input

                   Broca’s Aphasia
Lichtheim/Geschwind Model

                          Concepts    Association Cortex




                                       X
Ventral prefrontal                       Posterior Temporal
cortex Motor word                     Auditory word Cortex
        Comprehension    Arcuate      Comprehension
                         Fasciculus


   Speech motor output
                                              Auditory input

                 Wernicke’s Aphasia
Lichtheim/Geschwind Model

                          Concepts    Association Cortex




                         X
Ventral prefrontal                       Posterior Temporal
cortex Motor word                     Auditory word Cortex
         Comprehension   Arcuate      Comprehension
                         Fasciculus


   Speech motor output
                                              Auditory input

               Conduction Aphasia
More aphasias!
Aphasia type     Production      Comp          Repetition                 C
1. Broca’s      Non-fluent    Good            Poor                    5       4
                                                              1                           2
2. Wernicke’s   Fluent,       Poor            Poor
                paraphasic                                        M       3       A
3. Conduction   Fluent,       Good            Poor
                paraphasic
                                                              7                       8
4.Transcortical Fluent,       Poor            Good
sensory         paraphasic
5.Transcortical Terse,        Good            Good
motor           echolalic
6. Global       Poor          Poor            Poor

7. Aphemia      Dysarthric    Good            Limited only
                                              by dysarthria
8. Pure word    Normal        Very poor! As   Very poor
deafness                      if deaf!
Problems
• Models assumed to map directly onto
  underlying brain structures
• However, patients described as
  “Broca’s aphasics” had lesions in other
  areas
• Inadequate psychological concepts
  – e.g., “loss of motor images”
Broca’s Aphasia




• Damage to Broca’s area alone is not enough to produce
  Broca’s aphasia
• Usually involves Broca’s area + surrounding areas
  including M1 & insula.
Wernicke’s Aphasia




• Damage to Wernicke’s area alone is not enough to
  produce Wernicke’s aphasia
• Usually involves Wernicke’s area + surrounding areas
  including MTG & angular gyrus.
Conduction Aphasia




                            Common area?

• Damage to the arcuate fasciculus has not been associated
  with conduction aphasia
• Usually two lesion patterns: posterior STG (wernicke’s
  areas) and/or SMG
Transcortical Sensory Aphasia




                            Common area?

• Variable lesion patterns, mostly posterior to Wernicke’s
  area
• Deficit tends to be transient evolving into anomic aphasia
Transcortical Motor Aphasia




•Damage often anterior and/or superior to Broca’s area
Global Aphasia




• Tend to be large “peri-Sylvian” lesions
• But smaller lesions can also cause global aphasia
Aphemia




• Damage to lower motor cortex
Pure Word Deafness




• Damage to STG bilaterally is the most common pattern,
  although small left subcortical lesions have also been
  documented.
Neuropsychology Methods
• Early neuropsychology (Broca,
  Wernicke, Luria, et al.) made inferences
  based largely on associations
  – Identify a group of patients with fairly
    homogeneous functional impairment
  – fairly homogeneous physical impairment
  – Identify a “syndrome”
Problems


• Absence of rigid quantitative
  methodology of the day
• Patients not always carefully described
Resurgence of
          Neuropsychology
                in the 1960s
• Renewed interest in using
  neuropsychological cases to draw inferences
  about neural substrata of cognition
• More powerful inferential methods
• Emphasis on
  – Single case studies
  – Modern quantitative methods
Neuropsychology Methods
• Modern neuropsychology (e.g.,
  Warrington, McCarthy, Farah)
• Study single cases or groups with
  homogeneous physical impariment
• Focus on dissociations
Neuropsychology Methods
•   Single dissociation
•   Patient with lesion in brain region A
•   Performs well on task A
•   Performs poorly on task B
•   Inference: brain region A mediates
    performance on task B but not task A
Neuropsychology Methods
• Problems with single dissociations
• Does not rule out other interpretations
• Patient may suffer some global deficit (e.g.,
  general cognitive resource) that Task A does
  not require (e.g., because it is easier)
• Thus, performance differences between
  Tasks A and B may reflect task difficulty, not
  the function of Brain Region A
Neuropsychology Methods
• Double dissociation
• Patient with lesion in brain region A performs
  well on Task A and poorly on Task B
• Patient with lesion in brain region B performs
  well on Task B but poorly on Task A
• Inference: Brain region A mediates Task B
  Brain Region B mediates Task A
Neuropsychology Methods
• Since subjects are “equated” for
  damage, “global deficit” argument
  circumvented

• Tasks may be assumed to be modular,
  depending on separate neural
  processes or resources
Brain Imaging
       • We now have
         techniques that
         allow us to
         visualize the
         structure and
         function of the
         brain.
MRI
• MRI of a typical
  brain.
PET
 • Typical brain
   visualized with
   Positron Emission
   Tomography.
Functional
        Neuroimaging


                             -
                                 time (seconds)


             Deoxy
             Oxy




• Functional magnetic resonance imaging
Functional Neuroimaging

                      Passive word viewing
                      -Activates visual areas
                      -Particularly Left Occipital Cortex



                      Passive word listening
                      -Activates auditory areas
                      -Words (not nonwords)
                       particularly Left Temporal Cortex
                       (Wernicke’s area)
• Petersen et al., 1988 PET study
Functional Neuroimaging

                     Repeating words
                     -Bilateral activation of motor and
                     Sensory face areas
                     -No Activation of Broca’s or
                     Wernicke’s

                    Verb Generation
                    -e.g., Cake  “Eat”
                    - Left Frontal Lobe Activation
                     (Broca’s Area)
• Confirms neuropsychology research
What Have we Learned?
• What does localization tell us?
  – Where processing is occuring.
  – What is connected to what.
  – How many parts there are.
  – What happens when one part gets
    damaged.
• We’d like to know more.
What Would we Like to Know?
• How the parts work.
  – What are the rules and representations
    that are involved with language
    processing?
  – What is the nature of the processing that is
    taking place.
    • Digital/discrete/autonomous
    • Analog/continuous/integrated
ERP studies
• Semantics
  – N400
• Syntax
  – LAN
  – P600/SPS
N400 (2)
P600
Haagort et al. (2003)




Het vrouwtje veegde de vloer met een oude bezem gemaakt van twijgen (The
woman wiped the floor with an old broom made of twigs).
*Het vrouwtje veegde de vloer met een oude kliederde gemaakt van twijgen (The
woman wiped the floor with an old messed made of twigs).
P600 can serve as an index of
 frequency of a construction
• The captain liked the crew was unhappy
• The captain heard the crew was unhappy
• The captain believed the crew was unhappy
• A P600 was strongest in the first sentence, present in
  the second sentence, and not present in the third
  sentence.
• Larger P600 for syntactically complex sentences
  (Kaan et al., 2002).
Correct: De houthakker ontweek de schroef op dinsdag (The lumberjack dodged the
propellor on Tuesday).
Weak violation: *De houthakker ontweek de schroeft op dinsdag (The lumberjack
dodged the propelled on Tuesday).
Strong violation: *De houthakker ontweek de omdat op dinsdag (The lumberjack
dodged the because on Tuesday).
Combining methods
Summary
• The Wernicke-Geschwind model is
  being replaced by new models.
• New models focus on both neural basis
  and timing of language comprehension,
  and have moved beyond simplistic
  divisions of comprehension/production.

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Brain And Aphasia

  • 1. The Neuroscience of Language From Structure to Process
  • 2. Neuroscience and Language • Aphasia (or dysphasia) – Language disorder of auditory or oral speech, writing (agraphia), or reading (alexia) produced by injury to brain areas specialized for these functions
  • 4. Nature’s Experiments • Stroke – Status post 5 hours
  • 6. Blood-Brain Barrier • Blood is toxic to neurons. – Makes evolutionary sense. – Insulates brain from what’s in the blood.
  • 7. Paul Broca • Broca (1865) described patients who displayed halting, agrammatic speech – Content words were well preserved Paul Broca 1824-1880 – Function words (i.e., adjectives, articles) impaired
  • 8. Broca’s Aphasia • Patient “Tan” • Brain tumor in Left frontal brain region • Broca: Lesion disrupted speech
  • 9. Broca’s Aphasia • Broca’s Aphasia – Damage to “motor images” • Language comprehension skills relatively preserved • Typically observed in patients with damage to left inferior prefrontal cortex
  • 10. Broca’s Aphasia • “Yes… ah… Monday… er… Dad and Peter H… (patient’s name), and Dad… er… hospital… and ah… Wednesday… Wednesday, nine o’clock… and oh… Thursday… ten o’clock, ah doctors… two… an’ doctors… and er… teeth…yah Goodglass & Geschwind, 1976
  • 11. Carl Wernicke • Wernicke (1874) described patients whose speech is fluent, but has no informational value
  • 12. Wernicke’s Aphasia • Neologisms • Speech appears to have no information content • “fluent nonsense” • Preserved function words, impaired content words • Comprehension impaired • Even simple sentences not well understood • Associated with left temporal lobe damage
  • 13. Wernicke’s Aphasia • “Well this is… mother is away here working her work out o’here to get her better, but when she’s looking in the other part. One their small tile into her time here. She’s working another time…” Goodglass & Geschwind, 1976
  • 14. Articulatory Auditory speech speech “memories” “memories” Geschwind 1979 Scientific American
  • 15. Articulatory Auditory speech speech “memories” “memories” Production Comprehension problems problems + empty error-filled production
  • 16. Broca’s aphasia Wernicke’s aphasia Production Comprehension problems problems + empty error-filled production
  • 17. Lichtheim • Lichtheim (1885) – cases of patients able to understand and produce speech but unable to repeat words • “Conduction aphasia” • Hypothesis: Broca’s area (speech production) and Wernicke’s area (speech comprehension) intact
  • 18. Conduction aphasia Error-filled speech, but good comprehension Broca’s aphasia Wernicke’s aphasia Production Comprehension problems problems + empty error-filled production
  • 19. Broca’s, Wernicke’s Area, and Connections • Lichtheim’s (1885) and Geschwind’s (1965) model • Auditory input mediated by Wernicke’s area • Motor output mediated byBroca’s area • Regions connected by arcuate fasciculus
  • 20. Wernicke-Lichtheim “House” Model Concept representations C Widely distributed! M A Motor word images Auditory word images Broca’s area Wernicke’s area
  • 21. Lichtheim/Geschwind Model Association Cortex Concepts Ventral prefrontal Posterior Temporal cortex Motor word Auditory word Cortex Comprehension Arcuate Comprehension Fasciculus Speech motor output Auditory input
  • 22. Lichtheim/Geschwind Model Concepts Association Cortex X Ventral prefrontal Posterior Temporal cortex Motor word Auditory word Cortex Comprehension Arcuate Comprehension Fasciculus Speech motor output Auditory input Broca’s Aphasia
  • 23. Lichtheim/Geschwind Model Concepts Association Cortex X Ventral prefrontal Posterior Temporal cortex Motor word Auditory word Cortex Comprehension Arcuate Comprehension Fasciculus Speech motor output Auditory input Wernicke’s Aphasia
  • 24. Lichtheim/Geschwind Model Concepts Association Cortex X Ventral prefrontal Posterior Temporal cortex Motor word Auditory word Cortex Comprehension Arcuate Comprehension Fasciculus Speech motor output Auditory input Conduction Aphasia
  • 25. More aphasias! Aphasia type Production Comp Repetition C 1. Broca’s Non-fluent Good Poor 5 4 1 2 2. Wernicke’s Fluent, Poor Poor paraphasic M 3 A 3. Conduction Fluent, Good Poor paraphasic 7 8 4.Transcortical Fluent, Poor Good sensory paraphasic 5.Transcortical Terse, Good Good motor echolalic 6. Global Poor Poor Poor 7. Aphemia Dysarthric Good Limited only by dysarthria 8. Pure word Normal Very poor! As Very poor deafness if deaf!
  • 26. Problems • Models assumed to map directly onto underlying brain structures • However, patients described as “Broca’s aphasics” had lesions in other areas • Inadequate psychological concepts – e.g., “loss of motor images”
  • 27. Broca’s Aphasia • Damage to Broca’s area alone is not enough to produce Broca’s aphasia • Usually involves Broca’s area + surrounding areas including M1 & insula.
  • 28. Wernicke’s Aphasia • Damage to Wernicke’s area alone is not enough to produce Wernicke’s aphasia • Usually involves Wernicke’s area + surrounding areas including MTG & angular gyrus.
  • 29. Conduction Aphasia Common area? • Damage to the arcuate fasciculus has not been associated with conduction aphasia • Usually two lesion patterns: posterior STG (wernicke’s areas) and/or SMG
  • 30. Transcortical Sensory Aphasia Common area? • Variable lesion patterns, mostly posterior to Wernicke’s area • Deficit tends to be transient evolving into anomic aphasia
  • 31. Transcortical Motor Aphasia •Damage often anterior and/or superior to Broca’s area
  • 32. Global Aphasia • Tend to be large “peri-Sylvian” lesions • But smaller lesions can also cause global aphasia
  • 33. Aphemia • Damage to lower motor cortex
  • 34. Pure Word Deafness • Damage to STG bilaterally is the most common pattern, although small left subcortical lesions have also been documented.
  • 35. Neuropsychology Methods • Early neuropsychology (Broca, Wernicke, Luria, et al.) made inferences based largely on associations – Identify a group of patients with fairly homogeneous functional impairment – fairly homogeneous physical impairment – Identify a “syndrome”
  • 36. Problems • Absence of rigid quantitative methodology of the day • Patients not always carefully described
  • 37. Resurgence of Neuropsychology in the 1960s • Renewed interest in using neuropsychological cases to draw inferences about neural substrata of cognition • More powerful inferential methods • Emphasis on – Single case studies – Modern quantitative methods
  • 38. Neuropsychology Methods • Modern neuropsychology (e.g., Warrington, McCarthy, Farah) • Study single cases or groups with homogeneous physical impariment • Focus on dissociations
  • 39. Neuropsychology Methods • Single dissociation • Patient with lesion in brain region A • Performs well on task A • Performs poorly on task B • Inference: brain region A mediates performance on task B but not task A
  • 40. Neuropsychology Methods • Problems with single dissociations • Does not rule out other interpretations • Patient may suffer some global deficit (e.g., general cognitive resource) that Task A does not require (e.g., because it is easier) • Thus, performance differences between Tasks A and B may reflect task difficulty, not the function of Brain Region A
  • 41. Neuropsychology Methods • Double dissociation • Patient with lesion in brain region A performs well on Task A and poorly on Task B • Patient with lesion in brain region B performs well on Task B but poorly on Task A • Inference: Brain region A mediates Task B Brain Region B mediates Task A
  • 42. Neuropsychology Methods • Since subjects are “equated” for damage, “global deficit” argument circumvented • Tasks may be assumed to be modular, depending on separate neural processes or resources
  • 43. Brain Imaging • We now have techniques that allow us to visualize the structure and function of the brain.
  • 44. MRI • MRI of a typical brain.
  • 45. PET • Typical brain visualized with Positron Emission Tomography.
  • 46. Functional Neuroimaging - time (seconds) Deoxy Oxy • Functional magnetic resonance imaging
  • 47. Functional Neuroimaging Passive word viewing -Activates visual areas -Particularly Left Occipital Cortex Passive word listening -Activates auditory areas -Words (not nonwords) particularly Left Temporal Cortex (Wernicke’s area) • Petersen et al., 1988 PET study
  • 48. Functional Neuroimaging Repeating words -Bilateral activation of motor and Sensory face areas -No Activation of Broca’s or Wernicke’s Verb Generation -e.g., Cake  “Eat” - Left Frontal Lobe Activation (Broca’s Area) • Confirms neuropsychology research
  • 49. What Have we Learned? • What does localization tell us? – Where processing is occuring. – What is connected to what. – How many parts there are. – What happens when one part gets damaged. • We’d like to know more.
  • 50. What Would we Like to Know? • How the parts work. – What are the rules and representations that are involved with language processing? – What is the nature of the processing that is taking place. • Digital/discrete/autonomous • Analog/continuous/integrated
  • 51. ERP studies • Semantics – N400 • Syntax – LAN – P600/SPS
  • 53. P600
  • 54. Haagort et al. (2003) Het vrouwtje veegde de vloer met een oude bezem gemaakt van twijgen (The woman wiped the floor with an old broom made of twigs). *Het vrouwtje veegde de vloer met een oude kliederde gemaakt van twijgen (The woman wiped the floor with an old messed made of twigs).
  • 55. P600 can serve as an index of frequency of a construction • The captain liked the crew was unhappy • The captain heard the crew was unhappy • The captain believed the crew was unhappy • A P600 was strongest in the first sentence, present in the second sentence, and not present in the third sentence. • Larger P600 for syntactically complex sentences (Kaan et al., 2002).
  • 56. Correct: De houthakker ontweek de schroef op dinsdag (The lumberjack dodged the propellor on Tuesday). Weak violation: *De houthakker ontweek de schroeft op dinsdag (The lumberjack dodged the propelled on Tuesday). Strong violation: *De houthakker ontweek de omdat op dinsdag (The lumberjack dodged the because on Tuesday).
  • 58.
  • 59. Summary • The Wernicke-Geschwind model is being replaced by new models. • New models focus on both neural basis and timing of language comprehension, and have moved beyond simplistic divisions of comprehension/production.