Lec 1/ TB Ch 1, 2 Flashcards

1
Q
  • Reticulum theory
  • Cajal
    • silver nitrate method
    • 4 main discoveries
  • Brainbow method
  • The Brain Observatory
  • # neurons in brain
  • # neurons in cerebral cortex
A
  • Reticulum theory: all brain cells are fused together to form a big net or reticulum; this unit works in a holistic way
  • Cajal:
    • silver nitrate method: stain and drew what he saw from the microscope
      • Camillo Golgi invented the silver nitrate method
  • Santiago Cajal made 4 main discoveries
      1. Neurons only connect in a specific place called synapse
      1. Neurons are connected in principled (based on a set of rules), not randomly
      1. Electrical signals travel through neurons in only 1 direction; this allows systematic info flow thru circuits
    • (#4) neurons are independent units that are not joint together
  • Golgi supports the reticulum theory; against Cajal’s
  • Brainbow method: derived florescent proteins from glowing jellyfish to label specific neurons
    • It has 150+ colors
    • Cons: the color labelling is random
    • Current rs: try to tag different types of neurons w/ diff ranges of colors
      • Ex. red + orange for neuron type 1
      • Ex. Yellow + green for neuron type 2
      • Ex. Blue + purple for neuron type 3
  • Researchers attempt to disseminate results to broad audience
    • The Brain Observatory/ The brain’s Hubble telescope
      • Constructed the “Concise Digital Atlas of the Human Brain”
      • Ppl can inspect stained sections of the brain at various magnifications (ex. birds eye view of all structures, individual neurons)
  • Brains has 100 billion neurons, can stretch for 150,000 km
  • Cerebral cortex: 30 billion neurons, 1 million billion connections
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2
Q
  • LABEL THE DIAGRAM
  • Neuron fx
  • what are the fx of the following parts?
    • dendrites
    • axon
    • myeline sheath
    • nodes of ranvier
    • terminal buttons
    • synapse
  • Size of neuron in the retina?
  • Length of neuron spinal cord to toes?
  • 6 shapes of neurons
  • von Economo neurons
    • 2 places it is found
    • 7 fxs
    • Who has impaired von Economo neurons?
    • What animals have von Economo neurons?
A

Anatomy

  • Neurons’ function: signal transmission
  • Nucleus: Central region
  • Neurons look like trees: branching roots, long trunk, bushy crown
  • Dendrites: (tree in Greek) Branching roots, receive signals
  • Axon: (axle in Greek) long trunk, transmit signals
  • Myelin sheath: fat that give insulation to protect and accelerate signals
    • 1 mm long
  • Nodes of Ranvier: small spaces that separate the Myelin Sheath, rejuvenate (restore energy) the signals
  • Terminal buttons: the axon splits into segments, and it ends with the buttons, the buttons relay info to downstream neurons
  • Synapses: (syn = together; haptein = to clasp) tiny clefts b/w terminal buttons of message sending neurons and dendrites of message receiving neurons
  • Neurons vary in size and shape
    • Size: Neurons in the middle layer of the retina: 1 mm
      • Neurons that extend from spinal cord to toes: 1 m
    • Shapes: pyramidal, granular, stellate, chandelier, basket, fork
  • Some neurons are specialized for certain sensory or motor processing
  • Ex. von Economo neurons – found in a few brain regions (anterior cingulate, anterior insula); related to emotional processing, self-awareness, and social cog, interoception (can sense one’s internal bodily’s state), affect and communication in relationships
    • Those w/ mental disorders that impair mental capacities (ex. autism and schizo) have altered von Economo neurons
    • Socially complex mammals have many von Economo neurons (apes, elephants, whales, dolphins)
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3
Q
  • LABEL THE DIAGRAM
  • Process of how e- signals are conducted (3 steps)
  • Active conduction
A

Physiology

    1. Dendrites receives many inputs (in the form of e current) from other neurons
    1. E current moves to the cell body via passive conduction
      * Passive conduction: passive flow for e current from dendrite to cell body
    1. Current reaches the base (aka hillock of Axon hillock); if the total sum of the current exceeds the specific threshold, the neuron fires an action potential via active conduction
      * Action potential: aka spike, the neuron experience change in electrical property, this causes the net charge in the axon shift from -ve to +ve
      * Active conduction: reoccurring APs/spikes at the nodes of Ranvier, this allows the signal to travel along the axon w/o fading
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4
Q

Physiology cont

  • LABEL THE DIAGRAM
  • RMP?
  • Critical Threshold?
  • Process of AP: 3 steps
  • Fx of hyperpolarization
  • Signal speed of myelinated vs unmyelinated neuron
  • Fx of Nodes of Ranvier
  • Active conduction
  • Process to send signal across synapse
  • Summation effect
  • % of oxygenated blood used by brain
  • Size of arteries, arterioles, capillaries
A
  • By default, the neuron is more -ve in the inside (RMP: -70 mV)
  • When the input is +ve enough to increase the MP to -50 mV (aka critical threshold), this triggers an AP (happens in 1 ms)
      1. Na+ channels open; Na+ enters cell; cell depolarizes, the cell becomes more +ve than the external env
      1. Na+ channels close; K+ channels open (slowly K = couch potato); K+ leaves the cell; the cell becomes more -ve
      1. Since K+ channels close very slowly, the cell becomes hyperpolarized
        * Hyperpolarization: prevents another AP happen right away; ensures the signal travels forward, not backwards
  • Myelin Sheath: increase the speed an axon can transmit current
    • Unmyelinated: signal travels 1 m/s
    • Myelinated: signal travels 10 to 100 m/s
  • Nodes of Ranvier: small spaces b/w myeline sheaths; renews the AP perpetually
  • Active conduction: the cycle of Na+ influx and K+ outflow happens in each node; active conductions is the sequential (entire chain of) signal rejuvenation
  • Process to send signal across synapse
    1. When signals reach the terminal buttons, the have to cross the synaptic cleft (20 nm)
    1. Here, NT are released, and bind to specific receptors on the postsynaptic neuron
      * NT: chemicals that carry the signals across the synaptic cleft
      * If the presynaptic neuron is excitatory, the released NT make the postsynaptic neuron fire
      * If presynaptic neuron is inhibitory, the released NT make the postsynaptic neuron less likely to fire
  • Whether the target cell fires depend on the sum of numerous inputs
  • NOTE: the strength of individual inhibitory or excitatory connection is adjustable
    • Neuron activity uses a lot of energy
  • The human brain is 2-3% of the total body weight, but uses 20% of the oxygenated blood from the heart
  • Oxygenated blood from heart enters blood vessel network: arteries (5 millimetre) -> arterioles (30 micrometres) -> capillaries (10 micrometres; a bit wider than 1 RBC)
  • Capillaries
    • Have thin walls, oxygen can pass be extracted here
  • Veins: deoxygenated blood flows from brain to heart
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5
Q
  • What does neuron’s firing frequency indicate?
  • Why?
  • Why are assemblies of neurons arranged in multi-layered hierarchies?
  • Visual system
    • 2 main stages of processing
    • Describe Table detection network
  • How can Hierarchical coding networks be bottom-up feedforward?
  • How can Hierarchical coding networks have top-down feedback connections?
A

Representation

  • IOW, the neuron’s firing frequency indicates whether the preferred stimulus present
  • Neurons are tuned to respond to specific stimulus
  • Assemblies of neurons are put in multi-layered hierarchies; this arrangement help them capture complex patterns of info
    • Ex. Visual system - Object recognition
      • There are several stages of processing
        • Early stages: process elementary features of shape
        • Other stages: process more complex combination of the features
  • Ex. Table detection network
    • Level 1 (bottom): 1 cell code for “vertical edge”; 1 cell code for “horizontal edge”
    • Level 2: 1 cell code for length; 1 cell code for “corner”, and this cell receives input from the two cells in lv 1
    • Level 3: 1 cell code for legs; 1 cell code for rectangular surface, and it receives input (i.e. corners and lengths) from cells at lv 2
      • IOW, the cell can combine feature of corners and lengths to form the more complex “rectangular surface”
    • Level 4: 1 cell receives input from two cells in lv 3
      • IOW: this cell combines features of rectangular surface and legs to derive the final object representation – table
  • The table detection network resembles the hierarchical coding in the cerebral cortex, and other perceptual systems
      • Hierarchical coding networks are seen in bottom-up feedforward (use current feedback, and generate solutions for the future) connections
      • As a result, sensory signals can be matched w/ knowledge stored in LTM
  • Hierarchical coding networks are also seen in top-down feedback connections
    • This allows our internal priorities and predictions to guide our perceptual processes
    • Top-down connections go from central to peripheral brain regions
    • This allows us to reconstruct specific sensorimotor representations w/o external input
    • Ex. in our dreams, we conjure (bring forth) diff sights, sounds, and smells in our imagination
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6
Q
  • LABEL THE DIAGRAM
  • Planes
    • Sagittal
    • Coronal
    • Horizontal
  • Location of regions
    • Rostral (aka ?)
    • Caudal (aka ?)
    • Dorsal (aka?)
    • Ventral (aka ?)
    • Lateral
    • Medial
A

Navigating the Neural Landscape

  • Planes
    • Sagittal: section that separates left and right sides of the brain
    • Coronal: separate front from the back
    • Horizontal (aka axial/transverse): separates top from bottom
  • Locations of certain regions
    • Rostral/anterior: front of the brain (rostral; rooster; beak is in the front)
    • Caudal/posterior: back of the brain
    • Dorsal/superior: top of the brain
    • Ventral/inferior: bottom of the brain (V points down)
    • Lateral: outer left or right side of the brain
    • Medial: midline of the brain
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7
Q
  • LABEL THE DIAGRAM
  • Brainstem
    • 3 sections of brainstem
    • 2 things it maintains
    • Damage causes what?
    • Cranial nerves: what they do?
    • Where do cranial nerves the brain?
  • Thalamus
    • Where is it?
    • 3 fx
    • Which perceptual input does not pass through a specific nucleus in the thalamus?
    • Which nucleus signals from the retina enter?
    • Which nucleus signals from the cochlea?
    • 3 things thalamo-cortical loops facilitate?
A
  • Brainstem
  • has 3 main sections: medulla, pons, and midbrain
    • The 3 structures help maintain homeostasis and degree of wakefulness
    • Damage to the 3 structures -> coma or death
  • The portal (entrance) for cranial nerves
    • Cranial nerves receive sensory input from and send motor output to the head + neck
  • Thalamus
    • The large egg-shape structure on top of the brain stem
    • We have 2 thalamus, 1 for each hemisphere
    • (#3) It has many separate nuclei that direct the traffic in the brain
    • Thalamus is the “gateway to the cortex”,
        1. it relays info up to the cortex
          * All perceptual input (except smell) pass through a specific nucleus in the thalamus, then enters the appropriate cortical region
          * Ex. signals from the retina enters the lateral geniculate nucleus, then the primary visual cortex
          * Ex. Signals from the cochlea enters the medial geniculate nucleus, then the primary auditory cortex
          * Ex. signals from the basal ganglia, cerebellum, and amygdala enters the thalamic nuclei, then reach their cortical targets
        1. Thalamus receives feedback from the cortex
          * Each specific area in the cerebral cortex has bidirectional (2-way) connection w/ a specific part in the thalamus
          * These thalamo-cortical loops facilitate attention, STM, coordination of diff brain regions (ex. mental representations)
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8
Q
  • LABEL THE DIAGRAM
  • Hippocampus location
  • Fornix & Mammillary body fx
  • 4 steps Memory consolidation process
  • Patient HM
    • Which part was removed
    • What was the consequence of surgery
      • 3 things he can’t remember
      • 2 things he can
  • Amygdala location
    • 3 fx
    • Patient SM
      • 2 things she can’t do
      • 2 other specific fx of amygdala
A
  • Hippocampus
    • Aka Giant seahorse in Greek - Looks like a seahorse tail
    • Located in the temporal lobe
    • Fornix (blue box) & mammillary body: crucial to establish LT declarative memory
      • LT declarative memory: mem that can be verbally retrieved and reported
        • Ex. facts (current president)
        • Ex. autobiographical info (personal life info, where you went to HS)
    • Memory consolidation process
      1. hippocampus receives lots of convergent input from cerebral cortex
      1. Then it processes the input by registering the spatiotemporal info for related experiences
      1. Then, hippocampus retains these complex patterns
      1. After some time (ex. years), the hippocampus transfers this info back to the original cortical areas, which is stored semi-permanently in strongly weighted synaptic connections
      • Ex. Patient HM
        • Had amnesia after his medial temporal lobes were surgically removed
        • The medial temporal lobes were removed as a last resort to alleviate epileptic seizures
        • After the surgery, his seizures stopped, but
          • he can’t remember anything that happened to him since 16 yo
          • can’t remember simple things: who he had for his last meal, where he lived, own age
          • can’t learn the meanings of new words
        • IOW: this suggests hippocampus is essential for linguistic process
        • However, he has STM and can learn new motor skills
          • This suggest these abilities rely on other brain mechanisms
  • Amygdala
    • Located at the anterior tip of hippocampus
    • means almond in Latin
    • 1 Responsible for emo processing
    • 2 Helps us assess the significance/ value of the stimuli
    • 3 Sensitive to dangerous situations
    • Patient SM
      • Her amygdala developed abnormally in both hemispheres
      • 1 So, she cannot recognize fear expressions in other people’s faces
      • 2 She does not feel fear
      • Researchers brought her to haunted house
        • She reacted to the monsters by laughing, smiling, and talking to them
        • She scared the monsters too
      • She went through traumatic life events
        • She only felt angry and upset, never fearful
    • Amygdala helps promote adaptive b in threatening situations, and perceiving emotional prosodic patterns in speech (aka emotion in speech)
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9
Q
  • LABEL THE DIAGRAM
  • Basal Ganglia location
  • 4 main structures in Basal Ganglia
    • 2 structures in Striatum
    • Substantia nigra fx
  • How the Basal Ganglia works - 3 steps
  • 5 fx of basal ganglia
    • Cerebellum location
  • % of neurons in the brain in here
  • Purkinje neurons
  • 3 fx of cerebellum
  • If cerebellum is damaged, 3 main cons
A
  • Basal Ganglia
  • Basal ganglia: contains several integrated nuclei, located near the thalamus
  • Basal Ganglia contains the following structures
    • Striatum: includes the Caudate nucleus and Putamen
    • Globus pallidus or pallidum: has a lateral and medial portion
    • Subthalamic nucleus
    • Substantia nigra: located in the midbrain, generates dopamine, projected into the striatum
  • How the Basal Ganglia works
      1. Striatum receives signals from the cerebral cortex; Striatum recognizes. The signals from familiar situations (ex. I walk up to the door of my fav restaurant)
      1. Striatum sends this signal to 2 // routes
        * “go” pathway: this pathway implicitly learn what types of actions are adaptive in certain situations (ex. push the door)
        * “no-go” pathway: this pathway learns what types of actions are maladaptive in certain situations (ex. don’t pull the door)
      1. The output from these 2 pathways (i.e. recommendations on what to do or not) are sent to the frontal cortex via thalamus for further consideration
  • Basal Ganglia is essential for unconscious acquisition, selection, initiation, and cessation (aka habits OR procedural skills) in motor and cog control
    • It influences our motor control (ex. Parkinson’s, Huntington’s)
    • It influences our cog control (ex. schizo, OCD)
  • IOW: It influences overt b and covert thoughts
    • Cerebellum: located behind the medulla and pons
      • Aka little brain in Latin
      • But, it has more than two thirds of all the neurons in the brain
      • Contains Purkinje neurons
        • Purkinje neurons have many dendrites, and receive 200k signals from other cells
      • Cerebellum regulates muscle tone, balance, and movement control
      • Damage to cerebellum: leads to loss of balance, jerking/tremoring from arms and hands, impairs articulatory control causing dysarthria (slurred speech)
      • IOW: cerebellum influences movement and cognition
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10
Q
  • LABEL THE DIAGRAM
  • Cerebral cortex
  • Cortex in latin = ?
  • Size?
  • # of neurons?
  • # of connections
  • Grey matter - location?
  • White matter location?
  • 5 major lobes
    • 4 visible lobes
    • 1 invisible lobe
  • Sulci fx
  • Central sulcus
  • Lateral sulcus/sylvian fissure
  • 2 arbitrary lines
A

The Cerebral Cortex

  • Most highly evolved part of human brain
  • It takes up 3.5 pieces of letter sized paper; which is crumpled up to fit in the skull
    • The cortex has 30 billion neurons, each neuron contacts 1000 other cells
  • Gray matter: refers to the grey neuron cell bodies in the cortex
  • White matter: subcortical tissue is white due to myelinated axons

The Major Lobes–—Visible and Hidden

  • There are 5 lobes
  • 4 visible lobes on the cerebral cortex
    • Frontal
    • Parietal
    • Temporal
    • Occipital
  • 1 Invisible lobe: Insula
    • Insula = island in Latin
  • The borders are defined by sulci
  • Central sulcus: separates the frontal and parietal lobes
  • Lateral sulcus/sylvian fissure: separates the temporal lobes from the frontal + parietal lobes
  • An arbitrary line: separates the occipital lobe from the parietal and temporal lobe
    • This line connects the parieto-occipital sulcus (dorsal) with the preoccipital notch (aka dent) (located ventrolateral)
  • The other arbitrary line is perpendicular to the line above and extends to the end of the sylvian fissure
    • This separates the occipital from temporal lobe
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11
Q
  • LABEL DIAGRAM
  • Gyri
  • Sulci
  • 2 reasons why gyral-sulcal organization is adaptive
  • Axon density in Gyri vs sulci
  • How do sulci and gyri form?
  • shape and size ulci and gyri: env or genetic?
A

Gyral–Sulcal Organization

  • Gyri: bulges
  • Sulci: grooves
  • This gyral-sulcal organization is adaptive
      1. It squeezes a lot of surface area into a small space
        * Two thirds of the cortex is in the sulci
      1. It reduces the amount of axonal wiring
        * This reduces the distance signals travel
  • In humans and other mammals, axons projecting in the gyri are denser than axons projecting in the sulci
  • This suggest the gyral-sulcal patterns are caused by stiff axons constantly pushing harder against some regions on the cortical surface (like condom)
  • Since the skull is a physical constraint, the pushed regions protrude outward and form gyri; the other regions bend inward to form sulci
    • shape and size of gyri and sulci is mainly genetic
  • This is mainly genetically influenced (slightly env influenced)
  • Research showed that cortical convolutions are more similar in MZ than DZ twins
    • Diagram structures
  • Figure 1.17
    • 2 structure in the sylvian fissure on the dorsal surface of temporal lobe (aka supratemporal plane)
      1. HG = Heschl’s gyrus (aka transverse gyrus)
      1. PT = Planum temporale
    • Both are key for language and speech processing
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12
Q
  • cytoarchitectonic organization
  • cortex thickness
  • # of horizontal layers?
    • Layer I, II, III fx
    • Layer IV
    • Layer V
    • Layer VI
  • Define vertical columns
    • diameter?
    • # of neurons
  • Brodmann’s map
    • Which area does it map out?
    • # of Brodmann areas?
    • Areas range?
    • Why are some #s omitted?
    • 2 cons?
    • Which BA area is confirmed?
    • Which BA area is challenged?
    • How many distinct areas does each bran hemisphere has in reality?
    • Is Brodmann map a structural or functional map of the brain?
A

Cytoarchitectonic Organization

  • Cytoarchitectonic organization: whether there is packing density, and layers of diff types of cells
  • The cortex is 2-4 mm thick
    • Has 6 diff layers of cells stacked
    • Each layer of cells differs in morphological and connectional properties
      • Layers I, II, III: communicate w/ other cortical areas
      • Layer IV: receives input from thalamus
      • Layer V: sends output to subcortical motor structures
      • Layer VI: sends output to the thalamus
    • These layers are horizontal
    • There are vertical columns
    • Vertical columns: perpendicular to horizontal layers; basic functional units of the cortex
      • Each column is 0.4 mm in diameter
      • There are 100 neurons tuned to respond to similar features from the external or internal env
  • Scientists tried to make maps of the cortex
  • Brodmann’s map
    • the lateral and medial view of a human’s LH
    • there are 43 cortical areas OR Brodmann areas (Bas)
    • The areas ranged from 1 to 52
    • Brodmann omitted 12-16 and 48-51 b/c he used these #s for certain areas in other mammals, but he can’t find them in humans
  • In 1980s: BAs are often used w/ gyri and sulci labels to refer to specific cortical regions
  • It helped interpret fMRI
  • BAs weakness
      1. No info on the boundaries inside the sulci
      1. It is based on subjective observations; does not consider individual differences in anatomy
  • Modern imaging methods confirmed some BAs
    • Ex. BAs 44 and 45 do exist in the FL (BA 44 and 45 are usually collectively referred as the Broca’s area)
  • Modern imaging methods challenged some BAs
    • Ex. BAs 39 and 40 can be further carved into 7 areas
  • Currently, we know that each human brain hemisphere has 150-200 distinct areas
  • Main point: we should treat BA as structurally defined regions (rather than functionally)
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13
Q
  • LABEL DIAGRAM
  • Connectional Organization
    • What type of axons carry info b/w cortical regions?
    • Corpus callosum - what it does?
      • What matter type?
      • How many axons?
    • What do split brain patients help rs study?
    • What are Fasciculi?
    • Arcuate fasciculi fx
      • In which hemisphere?
      • which 3 regions does it secure?
      • What are the 3 connecting segments?
A

Connectional Organization

  • Cortical regions do not work in isolation; they work together to carry out complex mental processes (ex. language)
  • The signals are carried by axons that flow through white matter (like highways)
  • Corpus Callosum (CC): white matter tract w/ 100 million axons that connects the 2 hemispheres
    • The biggest and busiest tract
  • split-brain patients allow neuroscientists to examine the behavior from each hemisphere
  • Fasciculi: white matter tracts that connect diff cortical areas w/in the same hemisphere
    • Arcuate fasciculi: responsible for linguistics
      • In LH
      • It has 3 separate branches that undergird (secure) 3 regions
          1. Broca’s territory: posterior part of inferior frontal gyrus (BA 44 + 45) and adj portions of middle part of frontal gyrus and precentral gyri
          1. Wernicke’s territory: Posterior portions of the superior and middle temporal gyri
          1. Geschwind’s territory: supramarginal gyrus and angular gyri
      • The long direct segment: links Broca’s w/ Wernicke’s territory
      • Anterior indirect segment: links Broca w/ Geschwind’s territory
      • Posterior indirect segment: links Wernicke w/ Geschwind’s territory
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14
Q

Connectional Organization cont

  • LABEL THE DIAGRAM
  • 3 other language related fasciculi
  • Diffusion tractography
  • Human connector
A
  • Other language related fasciculi
  • Inferior fronto-occipital fasciculus: links inferior frontal area w/ occipital area
  • Inferior longitudinal fasciculus: link temporal areas w/ occipital areas
  • Uncinate fasciculus: link OBF area w/ anterior temporal areas
  • Main point: there are many long distance fibre tracts that tie the cortical regions together
  • This allows for producing and understand language
  • Diffusion tractography: measures passive movement of water along axons in white matter (uses MRI techniques)
  • This allows us to reconstruct the directions of fibre tracts (as seen in Fig 1.21, 1.22)
  • Human connectome: a project that maps all the connections in our brain
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15
Q
  • Sensory, Motor, and Higher-Order Systems
    • 3 primary sensory regions
    • Which cortex does visual info enters?
      • BA#?
      • Aka?
      • Location - which fissure?
      • Location - which part of which lobe?
A

Sensory, Motor, and Higher-Order Systems

  • Sensory: vision, audition, and somatosensory
  • Vision
    • Visual info enters the primary visual cortex in the occipital lobe
    • Primary visual cortex, aka BA17, V1, Striate cortex
    • This cortex is located in the calcarine fissure (Calc F), which is on the medial surface of the occipital lobe
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16
Q
  • Sensory, Motor, and Higher-Order Systems cont 2
    • Retinotopic organization
    • 2 separate streams at the anterior edge of occipital lobe
A
  • PVC has retinotopic organization
  • Retinotopic organization: preserves the spatial arrangement of signals from the retina
  • The PVC projects forward to other occipital areas that are specialized to extract info on specific attributes of visual stimuli (ex. form, color, motion, depth)
  • At the anterior edge of the occipital lobe, the flow splits into 2 separate streams
      1. “what” pathway: extend to ventral temporal lobe, it recognizes objects based on shapes, colors, and textures
      1. “where” path: extend to posterior parietal lobe, represent the locations of objects in relation to each other and in relation to the viewer’s body
        * Aka “how” pathway: promotes visuomotor transformations for object-directed actions
        • Ex. convert the position of a coffee mug based on the eye-centred coordinates into hand-centred coordinates

As a result, you can reach out and grasp the cup

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17
Q
  • LABEL THE DIAGRAM
  • Sensory, Motor, and Higher-Order Systems cont 3
    • Auditory
      • Location - gyrus?
      • Location - fissure?
      • Tonotopic organization
      • hemispheric asymmetries
      • Where are complex auditory fx done? - 3 parts
    • Somatosensory
      • postcentral gyrus fx
      • BA #s?
      • aka?
      • Define Somatotopic organization
        • large cortical areas means what?
      • 3 types of signals it processes
      • What info do signals from tendons provide?
      • 3 areas that do complex processing of somatosensory info?
A
  • Auditory
    • Primary auditory cortex is in the Heschl’s gyrus (located in the Sylvain fissure)
    • Primary auditory cortex area has a tonotopic organization
    • Tonotopic organization: cortical columns next to each other respond specifically to sound frequencies that are next to each other
    • PAC has hemispheric asymmetries: left PAC has more input from the right ear and vv
    • Complex auditory functions are done in the Planum temporale (PT), superior temporal gyrus, and middle temporal gyrus
  • Somatosensory cortex (aka postcentral gyrus)
    • postcentral gyrus (i.e. BA 1,2, and 3): Signals for felt shape and texture of objects, temp, pressure, and pain are first processed here
    • These BAs have diff fx; but are collectively referred to the primary somatosensory cortex (i.e. S1)
    • This region has rich somatotopic organization
    • Somatotopic organization: cortical representation that captures the body layout in a vertical way, upside down
      • w/ disproportionately large amounts of cortical territory for the most sensitive parts of the body (i.e. hands, feet, lips, tongue, and genitals)
      • NOTE: there is some discontinuities
        • Ex. face area is not next to the neck area; it is below the hand area
        • Ex. Genital area is not next to the upper leg area, but below the food area
    • the primary somatosensory cortex processes singals from the skin, muscles and tendons
    • Signals from the tendons carry proprioceptive info about the relative positions of one’s body parts in space and about the forces acting on them
    • Sophisticated analyses are done by the posterior and inferior parietal lobe, and the insula
18
Q
  • Sensory, Motor, and Higher-Order Systems cont 4
    • Motor cortex
      • aka - gyrus?
      • aka - cortex?
      • aka - BA#?
      • BA# for higher lv motor programming
        • what are the 2 areas
        • There fx
      • 2 higher order networks
        • Executive/supervisory network fx
          • 2 areas it engages
        • Mentalizing network fx
          • 3 areas it engages
A
  • Motor cortex (aka precentral gyrus)
    • Postcentral gyrus (somatosensory cortex) matches the precentral gyrus (motor cortex)
    • The precentral gyrus is the primary motor cortex; correspond to BA4, aka M1
    • Many higher lv motor programming regions are in BA6
      • Premotor cortex is in the lateral BA6
      • Supplementary motor area is in the medial BA6
        • Premotor cortex: contributes to externally triggered actions (ex. slam on brakes when you see a red light)
        • Supplementary motor area: contributes to internally triggered actions (ex. I get up and decide to go for a drive)
  • Higher-order systems
    • They engage other networks in other lobes of the brain
    • There are 6 other cortical areas
    • Talk about 2
        1. Executive/supervisory network
          * Depends on regions in the lateral PFC (BA 9, 10, 44, 45, and 46) and inferior parietal cortex (BA 39, and 40)
          * They are used in psychologically demanding situations that needs reasoning, planning, troubleshooting, multitasking, overcoming habitual responses, and keeping info in activated state
        1. Mentalizing network
          * Supported by medial PFC (BA 9 and 10), posterior cingulate (BA 23 and 31) and temporoparietal junction (intersection of BA 22, 37, and 39)
          * They help in social interaction; it is engaged when one tries to understand the overt b of other people, and covert mental states (ex. beliefs and desires)
19
Q
  • taxi drivers has larger ??
  • musicians has more ?? matter in what areas?
  • phoneticians have more tissue in ??
    • what leads to more tissue?
    • Is this nature or nurture?
  • phoneticians have more ?? matter in what area?
    • How many HG do they have in LH?
    • Is this nature or nurture?
    • What does this suggest
A

Box 1.1: Born for Phonetics?

  • Being an expert is influenced by structural and functional changes in the brain
  • Ex. London taxi drivers have larger hippocampus
  • Ex. Musicians gave more grey matter in auditory, motor, and visuospatial areas
  • Phoneticians (language experts)
  • Exp: Golestani et al 2011
    • Analysed the neuroanatomy of
      • 17 ppl w/ formal training in phonetic transcription
      • 16 controls
    • Results
        1. Phoneticians had greater SA and volume in the pars opercularis (a part) of the left inferior frontal gyrus
          * Aka the posterior portion of Broca’s area, known for phonological processing
          * The more training the phoneticians had; the more tissue detected in this area
          * This suggests transcriptional experience result in neural plasticity
        1. Phoneticians had more grey matter in the Heschl’s gyrus bilaterally (in L and RS
          * They also had a split or duplicated Heschl’s gyrus in the LH
          * Unlike Broca’s area, this area is not influenced by the length of training the phoneticians had
          * This suggests genetic factors are in play
  • Since gyri pattern stabilized by 7 yo
  • This may make these ppl more likely to be phoneticians or work in areas using detailed auditory processing
  • IOW, they are born for phonetics
20
Q
  • Which hemisphere does more language processing?
  • Broca area
    • Location - gyrus?
    • pars ?? and pars ??
  • Wernicke
    • Location gyrus?
  • Arcuate fasciculus
A

Language-Related Regions: Broca’s Area, Wernicke’s Area, and Beyond

  • Neural system for core components of language is located in separate areas
  • Human brain has a strong left hemisphere dominance for language
  • It has 2 main regions in this network
    • Broca’s area: includes the posterior portion of the inferior frontal gyrus, specifically the pars opercularis (BA 44) ad pars triangularis (BA 45)
  • Wernicke’s area: includes the posterior third of the superior temporal gyrus; it extends to the adjacent temporal and parietal regions
  • These 2 regions form a simple network
    • Broca’s area: represent the “motor images” of words, it is key for speech production
    • Wernicke’s area: rep the “auditory images” of words, it is key for speech perception
    • The 2 regions communicate via the arcuate fasciculus
  • Complex linguistic processes are done by cortical areas working together
21
Q

Neuropsychology

  • What do rs study in the field to understand the normal system?
  • 2 aims of neuropsychology
  • 19 C theory of the brain
  • equipotential
  • How is it disputed?
  • 2 types of dissociation
    • single dissociation
      • 2 alternative explanations
      • 2 solutions
    • double dissociation
    • Describe Ex. Linguistic distinction b/w nouns and verbs for single and double dissociation
A
  • Researchers study patients with brain damage to make new discoveries on the normal system’s design
  • Neuropsychology investigations have 2 general aims
      1. understand the cog architecture of language
      1. understand neural architecture

Single and Double Dissociations

  • 19C: ppl think brain operates as a unit
    • Equipotential: Each part contributes equally to every ability
  • If specific parts of the brain is damaged, specific abilities are impaired
    • Ex. damage Broca area → impair speech production
  • Dissociations are behavioural data
  • There are 2 main types
      1. Single dissociation
        * Patient is given 2 diff tasks and performs sig worse on X than Y
        * This infers that patient’s lesion has selectively disrupt some mental representations or computation that are needed by the poorly done task but not the better done one
        * Cons: this may not be an accurate conclusion; there are multiple explanations
        • In some situations, a single dissociation reflects specific mechanisms for 1 of the 2 tasks is disturbed
        • In other cases, this may reflect that the lesion disturbed the general processing resources that are needed for both tasks, and one of them needs more resources
        • Or the affected task may be harder
          * Solution
        • Ensure the tasks are matched on as many variables as possible (show similar accuracies, RT from healthy controls)
        • Double dissociation
      1. Double dissociation
        * Patient A does sig worse on task X than on task Y; Patient B does sig worse on task Y than on task X
        * The reduces the likelihood that patient’s performance is due to task difficulty
        * This suggests that each task needs some unique mental structure/operation and can be selectively disrupted
        * And the ability to do accomplish each task is impaired independently of the other
        * Double dissociation provides compelling evidence that the 2 tasks rely somewhat segregated cog mech
    • Ex. Linguistic distinction b/w nouns and verbs
      • Method: single dissociation - You gave patient 2-word retrieval tasks
          1. name pics w/ objects using the most appropriate noun
          1. name pics w/ actions using the most appropriate verb
      • Result: he did sig worse on object-naming task than action-naming task
      • Conclusion: You may conclude that his injury has compromised specific mechanisms essential for producing nouns but not verbs
      • May not be true
        • If the target nouns in object-naming task is way longer and less frequent than the target verbs in the action-naming task
        • Ex. orangutan vs walk
        • The patient may have more trouble accessing nouns than verbs as the object-naming tasking was more challenging
        • The patient’s injury then exacerbated his sensitive to the difference in difficulty
      • If you matched the 2 sets of target words for length and frequency, and showed most normal ppl produce both kinds of responses w/ similar speed and accuracy -> the evidence here better supports the conclusion
      • A better way is you demonstrate another patient who is tested w/ the same material showed the opposite dissociation
        • i.e. did sig worse on the action naming task than object naming task
      • This is a double dissociation
      • You can support the argument that 2 patients’ impairments most likely affect non-overlapping cog mech
      • You can also support that patient A had an impaired mech essential for noun but not verb production; patient B = opp
22
Q
  • Neuropsychology cont 2
    • Matzig et al 2009
      • Methods
      • Results
    • Rapp and Caramazza 2002 - In-depth study of patient KSR
      • Method
      • Results
      • 2 explanations
    • Groups and indiv
      • Group: pros, 3 cons
      • Syndrome
      • Indiv
A
  • Matzig et al 2009
    • 63 patients w/ noun-verb dissociations
    • The diff b/w 2 categories of words = 30%
    • Findings suggest that the 2 abilities (i.e. name objects w/ nouns, name actions w/ verbs) depend on at least partially distinct cog mech that can be selectively disrupted
    • Note: noun-related and verb-related picture naming deficits can b0 due to problems at several different lv of representation and computation
      • Ex visual, conceptual, grammatical, and phonological
    • Need more careful investigation
  • Rapp and Caramazza 2002
    • In-depth study of patient KSR
    • There is double dissociation in KSR who has brain damage
    • There is an interaction b/w v of grammar category and output modality
      • Patient had sig more trouble w/ nouns than verbs in oral production BUT more trouble w/ verbs than nouns in written production
    • Method: sentence completion paradigm w/ noun-verb homophones (same word diff meaning)
        1. Show patient KSR a “carrier sentence” & a pic of the object or action
          * Carrier sentence - Ex. Give me the (fish); I want to (fish)
    • Results: produced spoken nouns less accurately (40%) than spoken verbs (80%)
      • Produced written verbs less accurately (50%) than written nouns (90%)
      • 2 possible explanations
          1. The meanings of nouns and verbs are segregated w/in the semantic system
            * Although both types of concepts are still intact for KSR, she has 2 impairments
            * 1. Issue with the object concepts/nouns to speech (phonology)
            * 2. Issue with action concepts (verbs) to writing (orthography)
          1. The forms of nouns and verbs are separated in the speech and written output. KSR has 2 impairments
            * 1. Can’t form nouns in speech
            * 2. Can’t form verbs in writing
    • Main point: detailed neuropsychological studies uncover fine-grained disorders
  • NOTE: Single and double dissociations can apply to other domains in language (not just noun-verb tasks)

Groups and Individuals

  • Group studies and individual studies each have their own pros and cons
  • Group studies
    • Pros: allow rs to test hypothesis on correlations b/w disturbances from particular mental abilities and lesions
      • If a deficit reliably results from damaging a specific area, we should conduct group studies with
        • patients w/ + w/o impairment
        • patients w/ + w/o injury
    • Cons: limited on how carefully behavioural criteria is defined
      • Each patient has individual set of symptoms due to idiosyncratic brain damage
      • Group studies need to clearly specify the type of impairment that affects the behaviour to sort patients
    • Many rs focused on groups of aphasic patients who belong to certain “syndromes”
    • Syndrome: collection of symptoms that tend to co-occur statistically
    • Ex. Broca’s aphasia is a syndrome
      • Some Symptoms:
        • Can’t produce complex sentences
        • Worse retrieval of verbs than nouns
        • Poor articulations
    • Most rs treat patients as similar in their studies
    • Caramazza et al think this is invalid
      • These patients do not have all the same symptoms
      • They think syndromes like Broca’s aphasia lack empirical and theoretical use
    • Some think we should sort patients based on presence or absence of particular symptoms
    • Issue: symptoms like “worse retrieval of nouns than verbs in picture naming tasks” can have several different causes (see Rapp and Caramazza 2002 abv)
  • Individual studies
    • Provide insight on how the mind/brain is organized
    • Ex. Patient HM in Ch 1 helped rs develop theories on the role of hippocampus for LTM, and amygdala for emo processing
23
Q
  • Neuropsychology cont 3
  • MRI Basic mechanism
    • Brain is mainly made of?
    • 4 steps
    • Why can we see different type of organic tissues?
A

Visualizing the Brain: The Basic Mechanics of Magnetic Resonance Imaging

  • First technique: computed axial tomography (CAT/ CT)
  • Common technique: MRI
  • Obtains structural images of the brain in vivo
  • MRI Basic mechanism
    • Brain consists mainly of water; Each water molecule has 2 H atoms, and their protons are constantly spinning
    • This creates many tiny magnetic fields
      1. Normally, the orientation of the magnetic fields is random
      1. MRI machine changes the orientation by applying a strong magnetic field (1-10T) on the water molecules which are affected by gravity’s magnetic field (0.001 T). As a result, the protons align with this magnetic field.
        * Higher T/magnetic field, higher resolution
      1. Apply radio wave; this cause protons to absorb the wave energy and shift 90 deg
      1. Once the radio wave is turned off, the protons relax and align back with the external magnetic field. They emit a detectable signal.
    • We can see organic tissues (ex. white and grey matter) b/c they have diff densities of H atoms
    • This allows us to measure the differences in the relaxion rate of protons
24
Q
  • Neuropsychology cont 4
  • Types of brain damage
    • Stroke
      • Aka
      • what happens?
      • What is the most common type?
        • What happens to it?
      • 2 types of ischemia
        • differences?
      • What happens if it blocks brainstem?
      • What happens if it blocks white/cortical matter?
      • How to alleviate the deficits?
      • What happens if the deficit is not alleviated
    • Traumatic brain injury
      • Age group this is most common in?
      • 2 main types
      • When are they likely to happen?
      • Gabrielle Giffords
    • Neurodegenerative and Infectious Diseases
      • How are these diseases different that stroke and TBI?
      • their 2 main targets
      • the 10 disorders related to language (7 affect language in indirect ways; 3 PPA)
        • disorder name
        • type
        • affected brain region
      • PPA?
      • voxel-based morphometry (VBM)
        • What imaging technique?
        • 2 step process
A

Types of Brain Damage

  • Stroke (aka cerebrovascular accident – CVA)
    • Occurs when blood supply to a particular part of the brain is interrupted
    • Common type: ischemia
      • Blood vessel in the brain is obstructed by a clot, and this deprives downstream tissue of oxygen
      • Thrombotic ischemic stroke: the clot forms w/in the blood vessel eventually gets clogged
      • Embolic ischemic stroke: clot originates in a diff part of the circulatory system (ex. heart, large vessels of upper neck and chest)
        • This travels up to the brain and get stuck in the capillaries
    • If it blocks sectors of the brainstem for basic life functions -> person will lose consciousness and die in minutes
    • If it blocks certain cortical and white matter areas that support specific cog capacities, the person lives but the cog capacities are compromised
      • Ex. strokes that block the posterior cerebral artery that supplies to occipital areas -> visual impairments
      • Ex. Strokes that block the middle cerebral artery that supplies to the perisylvian areas for linguistics -> linguistic impairments
    • Deficits can be partially or completely alleviated if blood flow is rapidly restored to the deprived tissue
    • If not, the tissue will die and become absorbed
    • This leaves a cavity that gets filled w/ CSF
    • Traumatic Brain Injury
      * The most frequent form of brain damage suffered by ppl under 40
      * 2 main types
      * 1. Closed head injury
      * Occurs when person sustains a sudden blow to the head that damages the brain and leaves the skull intact
      * Ex. car accidents, sporting events
      * Ex.
      * Superficial damage to left perisylvian territory
      * Patient had aphasia, but not motor or sensory deficits
      * 2. Open head injury
      * Both the brain and the skull are penetrated by an object (ex. bullet, shrapnel)
      * Ex. Gabrielle Giffords
      * Forced to resign from her set in US House of Representatives after being shot in the head on 2011
      * Remains seriously disabled
      • Neurodegenerative and Infectious Diseases
        • Brain damage from stroke and TBI have immediate effects
        • Neurodegenerative and infectious disease are progressive
        • There are many disorders and all involve gradual atrophy (i.e. tissue loss) in specific regions/sets of regions in the brain
        • These disorders target particular networks that
          • Have a strong functional-anatomical connection
          • Are sig correlated to grey matter volume
        • 10 disorders have cons for language (table)
          • First 7 affect language in indirect ways
            • They impair certain aspects of perception, memory, cognition, and motor control that are necessary for efficient production and comprehending words/sentences
          • Last 3 are PPA
            • PPA = primary progressive aphasia
            • PPA target core components of language circuitry
            • Distinctive patterns of slowly worsening language deficits
            • Distinctive patterns of regionally specific atrophy
            • The 3 PPA have nonoverlapping distributions of cortical atrophy
            • 3 types
              • Nonfluent/agrammatic variant: tissue loss in left inferior frontal cortex
              • Semantic variant: anterior temp lobe (bilaterally)
              • Logopenic variant: left temperoparietal area
        • voxel-based morphometry (VBM)
          • A type of MRI
            1. It divides the brain into thousands of voxels (i.e. tiny cubes)
              * Each cube is a volume (similar to pixels)
              * 2. determines concentration of grey and white matter in each voxel
              * Rs can determine the locations and magnitudes of atrophy in patients w/ neurodegenerative/infectious diseases
              * Then compare them to healthy controls
25
Q
  • Neuropsychology cont 5
  • Tumours
    • Definition
    • Glioma: definition
    • Growth rate
    • Kinno et al 2009
      • Japanese-speaking patients w/ gliomas affecting Broca’s area
      • Method
      • Results
  • Using behavioral data and lesion data
    • 3 steps Lesion Overlap and Subtraction Analysis
    • Tranel and Kemmerer 2004 - understand neural basis of impaired knowledge on meanings of location prepositions
      • Locative prepositions
      • Part 1
        • Methods
          • 4 tasks
        • Results
      • Part 2
        • Method
        • Results
    • Voxel-Based Lesion–Symptom Mapping (VLSM)
      • 3 steps
      • Wu et al 2007 - VLSM & locative prepositions
    • 3 caveats About Neuropsychological Research on Structure–Function Relationships
    • 2 ways where areas are dysfunctional and may not show up on MRI
    • Apraxia of Speech (AOS)
    • Nina Dronkers 1996
      • Method
      • Results
      • 3 limitations
    • Argye Hillis 2004
      • Methods
      • Results
    • What went “wrong” w/ Dronker’s study?
A
  • Tumours
    • Lesions can be due to tumors
    • Tumours: masses of tissue that grow abnormally and serve no physiological purpose
    • Several type (classified based on how they dev, and likelihood to recur after surgical removal)
      1. Glioma
        * Come from white matter and expand outward -> destroy or displace neurons
        * Variable rate of growth (can be v slow or fast)
        * Glioma patient studies are useful
        * Kinno et al 2009
        * A group of Japanese-speaking patients w/ gliomas affecting Broca’s area had sig worse comprehension for passive sentences than active ones
        • Ex. Passive – Person A is being pushed by Person B
        • Ex. Active – Person A is pushing Person B
        • Original stimuli: Japanese sentences paired w/ scenes with 2 ppl
          * Another study
        • Showed these patients’ lesion sites overlapped w/ an area that was engaged sig more by passive than active sentences based on another fMRI study
        • They compared them to healthy controls who did the same task
          * Since the neuropsychological data and fMRI data aligns, this shows reliable inferences can be drawn from Glioma studies

Relationships Between Behavioral Data and Lesion Data

  • -> discuss how groups studies are conducted
  • Lesion Overlap and Subtraction Analysis
      1. Two groups of patients
        * Gp A w/ deficit of interest
        * Gp B do not have it
      1. Each groups has separate procedures
        * Reconstruct the patients’ lesions on a standard brain template
        * Calculate how much overlap for every voxel
      1. Lesion overlap map for patients w/ deficit minus lesion overlap map for patients w/o deficit -> remaining area = areas of damage that is linked w/ deficit
    • Tranel and Kemmerer 2004
      • Tried to identify the neural basis of impaired knowledge on meanings of location prepositions
      • Locative prepositions:
      • Ex. in, on, around, through, above, below
      • Methods
        • 4 tasks given to 78 brain-damaged patients (from stroke)
          1. Naming (80 items)
            * Ppl are shown objects and are asked to orally name the location of one object relative to another
          1. Matching (50 item)
            * Ppl are shown 3 spatial arrays of objects w/ a preposition
            * Told to choose which array best represent the preposition
          1. Odd one out (45 items)
            * Ppl are asked to choose which one involves a type of relationship that is different from the other two
          1. Verification (44 items)
            * Ppl are shown a spatial array of abstract shapes together and a preposition
            * They need to decide whether the preposition correctly describes the array
            * * Results
        • Very variable: some failed none, some failed one task, some failed all 4 tasks
        • Those who failed all 4 tasks may have impaired knowledge on the location preposition meanings
        • Those who only failed only 1 task had unimpaired knowledge on meanings but may have individual disturbances on certain processes
      • Part 2
        • Rs created lesion overlap maps for those w/o impairments; and one for those w/ impairments
        • Then they subtract the maps from each other
        • Result shows defective knowledge on the meanings of location prepositions is associated w/ damage in some areas in LH
          • The cortex and underlying white matter in the inferior frontal, inferior parietal, and posterior superior temporal regions (red, orange, yellows)
            • The analysis includes patents w/ the deficit of interest and those w/o
      • So, the results provide strong evidence that the lesions mentioned are more likely than not to cause the deficit
  • Voxel-Based Lesion–Symptom Mapping (VLSM)
      1. Administer a task to patients w/ various lesions
      1. Patients’ lesion sites are transferred to a standard brain template
        * For each voxel, 2 gps of patients are formed:
        • Those w/ brain lesions vs those w/o
      1. For each voxel, rs do a t-test to compare the behaviour performances of patients in the lesioned and non-lesioned groups
        * The resulting t-values measures the degree on whether decreasing task performance is related to the presence vs absence of certain brain areas being damaged
    • IOW, VLSM allows rs to identify the neural reasons of deficits, quantify brain-behaviour relationships w/o classifying patients based on their impairment
    • Wu et al 2007
      • Used VLSM to study impaired knowledge of meaning of location prepositions
      • 14 patients w/ LH lesions
      • They do several tasks the sentence w/ location prepositions w/ 1 of 4 pics
      • Results: accuracy range b/w 43% to 100%’ mean = 86%
      • The lesioned areas are consistent w/ those from Tranel and Kemmerer (2004 – abv)
  • A Few Caveats About Neuropsychological Research on Structure–Function Relationships
    • 3 caveats
      1. To infer a causal relationship b/w an impaired ability and damage to a specific region, need to show that
        * Patients w/ deficit have lesions at that site
        * Patients w/ lesions at that site tend to have the deficit
      1. Deficit-lesion correlations for those w/ stroke/TBI in the chronic period can be diff from those in the acute period
        * Chronic period: 6+ mo after lesion onset
        * Acute period: less than 6 mo after lesion onset
        * This is b/c acute patients w/ small lesions recover fast; the intact areas “take over” the affected fx
      1. Some impairments may be due to areas that are not structurally damaged
        * These areas are dysfunctional and may not show up on MRI
        * 2 ways
        * A. Hypoperfusion: receives enough blood supply to survive but not enough to operate normally
        * B. Diaschisis: they depend on axon input from the site of structural damage; but the input is no longer available
    • Apraxia of Speech (AOS): disorder of articulatory programming in Broca’s aphasia
      • AOS affect high-level motor aspects of orchestrating speech
      • This leads to distorted consonants, vowels, and prosody
      • Patients know what they want to say and how it should sound, but cannot coordinate the articulators (i.e. lips, tongue, jaw, and palate) to produce the output
    • Nina Dronkers 1996
      • Showed 25 chronic stroke patients w/ AOS had 100% lesion overlap in one LH region – superior anterior part of the insula
      • 19 chronic stroke patients w/ AOS had that damage
      • 3 limitations (related to 3 caveats above)
          1. Results show that AOS is associated w/ a damaged area of insula; but it doesn’t tell if damage to that area give rise to AOS
            * It is possible that AOS may also involve damage in other areas
          1. All the patients were chronic; acute patients may show diff deficit-lesion correlation
          1. The study only focused on structural damage
            * Other areas that are intact but are dysfunctional by contribute to AOS
    • Argye Hillis 2004
      • Addressed the 3 limitations in Dronker’s study
      • Methods
          1. Selected 80 stroke patients based on site of structural damage
            * 40: lesion on any part of the left insula
            * 40: spared that area
          1. Patients were eval for AOS when they were acute (24 hr of stroke onset)
          1. Patients were scanned of tissue damage, and also areas of hypoperfusion (sig reduced blood flow)
      • Results
        • Abnormalities in region of interest (superior anterior part of left insula) is not sig related to AOS
          • 12 out of 29 patients (40%) who had damage or hypoperfusion in that area showed AOS
          • 19 out of 51 patients (40%) who did NOT hv damage or hypoperfusion in that area showed AOS
        • AOS is actually associated w/ abnormalities in Broca’s area
          • 26/30 (90%) who had damage or hypoperfusion in that area showed AOS
          • 5/50 (10%) who did NOT hv damage or hypoperfusion in that area showed AOS
      • What went “wrong” w/ Dronker’s study?
        • The insula is the most vulnerable to stroke due to anatomy
        • Maybe 80% of Dronker’s chronic patients had damage to Broca’s area, while the remaining patients had hypoperfusion in that area
26
Q

Functional Neuroimaging

  • aka
  • Vampire theory
  • Fulton 1930 & Walker who had many congenitally abnormal blood vessels in occipital cortex
  • Positron Emission Tomography (PET)
    • fx
    • process
    • spatial resolution & temporal resolution
A

Functional Neuroimaging

  • Aka hemodynamic methods (b/c vampire theory)
    • PET & fMRI (more common)
    • Map the function in human brain
    • Vampire theory: the more active regions suck more blood
    • Key process
        1. When the cog task uses a particular brain area, the active neurons in that area rapidly consume the oxygen available in the local capillaries
        1. A few seconds later, there is an overabundance of oxygenated blood
  • Fulton 1930s
    • Dr. Fulton worked w/ Walter who had many congenitally abnormal blood vessels (arteriovenous malformation) in occipital cortex
    • It causes headaches and visual disturbances
    • When blood pass through those vessels intensely, it produces a pulsing sound that the patient hears as a humming sound
    • Dr. Fulton can also hear it when putting a stethoscope at the back of patient’s head
    • Dr. Fulton found that the rushing sound is correlated w/ Walter’s heartbeats and visual experiences
    • Ex. If Walter lied in the dark for a few minutes and suddenly use his eyes OR reading smth attentively, the sounds gets louder
    • This shows that changes in blood flow reflect changes in neural activity in certain regions, and this affect cog changes

Two Techniques

  • Positron Emission Tomography (PET)
    • Measures regional cerebral blood flow, which infers regional neural activity
    • The machines tracks the distribution of radioactive isotopes throughout the brain
      1. Rs create an isotope in a medical cyclotron, and place it in water
      1. This radioactive water is sent to the imaging facility, and rs inject it into the subject’s bloodstream. The bloodstream carries the radioactive water into the subject’s brain.
        * 15 O is the most common isotope used
        * It has 8 p+ and 7 n0
        * When it decays, it releases a proton that is +vely charged (i.e. positron)
        * The positron travels and is attracted to surrounding/ambient electron; this causes the electron and positron to be destroyed (annihilated)
        * As a result, 2 photons are released in opposite directions
        * These photons exit the head and are picked up by the rings of detectors in the PET scanner
        * The computer reconstructs the annihilation location in the subject’s brain
        * If there is a large # of annihilation events in an area, this suggests there is more blood flow in that area. This in turn indicates there is more neural activity
    • Spatial resolution: 10mm (kinda bad)
    • Temporal resolution: 30s (bad)
      • Reason: need to average the data over the length of time to get a good signal-to-noise ratio??
27
Q

Fx neuroimaging cont 2

  • fMRI
    • What is it sensitive to?
    • BOLD
    • 12 second hemodynamic response function (4 steps)
    • Is BOLD correlated to input or output processing of neurons
    • Spatial and temporal resolution
    • 3 Pros of fMRI over PET
    • 2 Cons
A
  • Functional Magnetic Resonance Imaging (fMRI)
    • Sensitive to degree of oxygenation of blood in diff parts of brain
    • Blood oxygenation level dependent signal (BOLD)
      • Endogenous contrast agent: deoxygenated blood reduces the MRI signal; oxygenated blood increases this
      • When hemoglobin is not carrying any oxygen, it behaves like a small magnet and disrupts the local magnetic field
      • This reduces the signal received by the fMRI machine
      • When hemoglobin is carrying oxygen, it is less magnetic
      • The fMRI can still detect them as a small increase in resonance signal
    • 12 second hemodynamic response function
        1. When neurons in a specific area increase in activity, they consume the oxygen that is immediately available
        1. As a result, there is more deoxygenated blood in that area, and the fMRI will pick up a small decrease in BOLD signal for 2s
        1. In the next 5s, more oxygenated blood is detected in that area, and the fMRI detects this as a gradual increase in the BOLD signal
        1. For the next 5s, the signal will return to baseline and briefly undershoots
    • The BOLD signal maybe more correlated to the input processing of neurons rather than output
    • Spatial resolution: 1-3 mm
    • Temporal resolution 50-100 ms
    • Pros of fMRI over PET
        1. Cheaper: doesn’t need a medical cyclotron
        1. Safer: doesn’t use radioactive isotopes
        1. Better spatial and temporal resolution
    • Cons
        1. It is very noisy
        1. BOLD signals are distorted near air-filled cavities
          * It is tricky it get data from some brain regions, esp the anterior temporal lobes located near the sinuses
          * (rs found a way around this)
28
Q

Fx neuroimaging cont 3

  • Standardized Three-Dimensional Coordinates for Defining Stereotaxic Brain Space
    • Units?
    • What do we do in fMRI studies
    • Origin?
    • x axis plane?
    • y axis plane?
    • z axis plane?
    • 2 diff atlases/3D grids
    • Fedorenko and Kanwisher 2009
      • Cons of analysing multiple subjects for fMRI
      • Solution
        • 3 steps
A

Standardized Three-Dimensional Coordinates for Defining Stereotaxic Brain Space

  • In fMRI studies, rs transform the anatomic confiscation of each subject to a standard brain template, then they combine data across subjects
  • Stat sig activations are identified and reported in the common stereotaxic space of the standard brain template
  • The brain template uses the x,y,z coordinate system
  • The origin is the anterior commissure
    • X-axis: right-left dimension (right = +ve; left = -ve)
      • Y-axis: anterior-posterior (anterior = +ve; posterior = -ve)
      • Z-axis: superior-inferior (superior = +ve; inferior = -ve)
  • 2 diff atlases/3D grids (unit: voxels)
      1. Talairach and Tournous
        * Based on anatomical data from 1 post-mortem brain
      1. Montreal neurological institute (MNI) 1994
        * Based on MRI scans from 305 healthy subjects
  • They are similar but not identical
  • Old studies voxel size: 3-5 mm
  • New studies: 1 mm
  • Potential cons of analysing multiple subjects for fMRI
    • Fedorenko and Kanwisher 2009
      • This may obscure indiv diff in neural organization of language by removing anatomical variability b/w ppl’s brains
      • Potential solution: Plot each subject’s unique functional data on his/her anatomical data
          1. Locate a region of interest (ROI) in each subject
            * Ex. Fusiform Face Area – sensitive to seeing faces
          1. Test for other response properties for that ROI
            * Ex. See if FFA is more sensitive to upright vs inverted faces
          1. Conduct analyses across subjects using data from corresponding functional regions rather than drawing from the same locations in Talairach/NMI space
      • May provide more insight
29
Q

Fx neuroimaging cont 4

  • Blocked Versus Event-Related Designs
  • Block design
    • When do we use it? (neuroimaging technique)
    • fx
    • How it works
  • fMRI study design options
    • block design
      • 2 differences compared to other techniques when using block design
    • event-related design
      • definition
      • how it works
      • 3 pros
A

Blocked Versus Event-Related Designs

  • How should we order individual trials in experimental condition?
  • Ex. You want to identify and compare brain regions that respond to words for animals (ex. rabbit) and words for tools (ex. pencil, knife)
  • -> How do we sequence the trials for animal words and tool words?
  • This depends on the imaging technique
  • PET
    • Need to use block design b/c PET has poor temporal resolution
    • Block design
      • Trials for the same experimental condition are grouped together in blocks
        • Here, the subjects must do the same type of task for the same type of stimuli throughout the block
      • Ex. You set up 3 blocks in your PET study (each block = 1 min)
        • Block 1: Condition A – ppl read words for animals
        • Block 2: Baseline condition – ppl lie in the scanner; do nothing
        • Block 3: Condition B – ppl read words for tools
      • These blocks are separated by 10 min periods – this is to allow time for the radioactive isotope subjects receive at the beginning of each block to decay
      • Hypothetical results:
        • Strongest signal: Block 1/Condition A- read words for animals
        • Weakest signal: Block 2/Baseline – rest
        • In b/w: Block 3/Condition B – read words for tools
        • fMRI
    • Since it has better temporal resolution, you have more design options (2 options)
    • Option 1: You can use block design, but there will be
      • More blocks per condition
      • Shorter blocks (30s)
      • Same hypothetical results:
        • Strongest signal: Block 1/Condition A- read words for animals
        • Weakest signal: Block 2/Baseline – rest
        • In b/w: Block 3/Condition B – read words for tools
        • Option 2: Event-related design
      • The trials belonging to diff experimental conditions are randomly scattered
      • Each trial has its own hemodynamic response
      • These indiv signals are later extracted and averaged based on the condition
      • Same hypothetical results
        • Strongest signal: Condition A- read words for animals
        • Weaker singal: Condition B – read words for tools
          • Why are event-related design popular?
      1. Allow rs to randomize stimuli
        * Reduces the chance the results are influenced by habituation, anticipation, or strategic processing (which r more likely in block design)
      1. There’s more experimental flexibility than block design
        * Trials can be sorted by diff criteria
        • Ex. conditions, uncontrollable factors (trials subjects responded incorrect)
      1. Can see more details on amplitudes and time trends of hemodynamic response in diff brain regions
30
Q

Experimental paradigms

  • Which study design are experimental paradigms usually used w/?
  • Subtraction paradigm
    • What study designs can used this paradigm?
    • What neuroimaging methods can use this paradigm?
    • Methods - how to use
    • Narain et al 2003: Aims to disclose brain regions linked w/ auditory processing of intelligible vs unintelligible sentences
    • 4 conditions
    • Methods
    • Final activation map
  • Correlation paradigm
    • What study designs can used this paradigm?
    • How it works
    • Mobbs et al 2010 - Looked at the neural activity associated w/ monitoring the threat value of a tarantula
      • Methods
      • Results
    • Davis and Johnsrude 2003 - Used correlation paradigm to further explain the neural underpinnings of intelligible and unintelligible
      • Methods
      • 5 conditions
      • 3 diff degrees of intelligibility
      • Results
        • Subtraction paradigm
        • Correlation paradigm
        • Follow up analyses
    • Multivariate Pattern Analysis
      • How is it different from subtraction and correlation paradigm?
      • What it does?
      • Mur et al 2009/ Raizada et al 2010
        • orthogonal pattern detection
      • Abrams et al 2013 -Used MVPA to examine the neural substrates that process intelligible vs unintelligible utterances w/ better precision
        • Methods
        • Analysis
          • subtraction
          • MVPA
        • Results
A

Some Basic Experimental Paradigms

  • Subtraction, correlation, and pattern analysis are mainly used w/ event-related design, but occasionally w/ block design
  • Subtraction paradigm
    • First paradigm used in fx neuroimaging
    • Can be used with blocked and event-related designs
    • Used by fMRI and PET rs
    • Helps rs isolate neural correlates of specific cog capacities
      • Study design: 2 conditions
        • Experimental: ability of interest
        • Baseline/Control: does not include ability of interest, equivalent in other aspects
      1. Collect imaging data
      1. Map of brain activity w/ experimental condition “-“ map of brain activity w/ control condition
        * The regions that still show up is sig engaged in the experimental condition; this is treated as uniquely contributing to the ability of interest
    • To enhance reliability, we include
      • multiple experimental conditions that require the ability of interest (Ex. conditions A and B)
      • multiple control conditions that do not (Ex. C and D)
    • This allows rs to conduct conjunction analyses that increase the likelihood of pinpointing brain regions responsible for the interested ability
      • Ex. If each experimental condition is well-matched w/ a particular control condition
        • Ex. A w/ C, B w/ D
          1. Perform separate subtractions b/w corresponding conditions
          1. Conjoin outcomes of the independent analyses [(A – C) + (B – D)]
        • The resulting brain map is quite restricted, and helps narrow the neural underpinnings of specific cog capacities
    • Narain et al 2003
      • Aims to disclose brain regions linked w/ auditory processing of intelligible vs unintelligible sentences
      • 4 conditions: 2 using intelligible stimuli, 2 using unintelligible stimuli
      • Intelligible conditions
        • A. Speech (Sp): normal sentences
        • B. Vocoded speech (VCo): sentences that are artificially altered to have a rough sound quality but still can be understood
      • Unintelligible conditions
        • C. Rotated speech (RSp): Sentences that are spectrally rotated (i.e. inverted) so they sound like “alien” language;
          • but still have phonetic features (ex. voiceless fricatives – “f”, “th”)
        • D. Rotated vocoded speech (RVCo): Sentences that have been vocoded and spectrally rotated (i.e. inverted) so they sound like intermittent fluctuating static
      • 11 ppl listened to blocks of stimuli from each condition during fMRI scanning
      • All state they can only understand stimuli 2 out of 4 conditions
      • Rs identified the brain regions that mediate extracting meaning from intelligible utterances using the “subtraction” method
        • [(A – C) + (B – D)]
        • [(Sp – RSp) + (VCo – RVCo)]
          1. Do 2 separate subtractions
            * Each involves contrasting the activation evoked by intelligible stimuli against the matched set of unintelligible stimuli
            * Intelligible and unintelligible conditions differ only by spectral rotation
          1. Conjoin (add) activation maps from the 2 separate subtractions; this identifies the commonality
            * This captures brain areas that respond to both intelligible stimuli
            * This rule out areas that respond to only one intelligible stimuli and areas that respond to both unintelligible stimuli
      • Final activation map
        • Only 3 clusters of voxels are stat sig
        • They are in the left lateral temporal lobe
        • Some portion overlap with Wernicke’s area,
        • Damage to that area disrupts phonological and lexical aspects of spoken language comprehension
        • It’s possible lateral temporal region contributes to high-level auditory sentence processing, not low-level acoustic and phonetic info
      • Note: analysis was restrictive
  • Correlation paradigm (aka parametric paradigm)
    • Compatible w/ blocked and event-related design
    • The ability of interest here is a continuous variable that can vary; it is not all-or-nothing
    • Subjects perform a series of tasks that uses the ability of interest in diff extent
      • Rs look for brain regions where the activity level changes correspondingly
    • Mobbs et al 2010
      • Looked at the neural activity associated w/ monitoring the threat value of a tarantula
        1. Ppl were placed in fMRI, one foot is in a box w/ 5 compartments
        1. Ppl think they were seeing a live camera feed where a rs move a large tarantula from one box to another, closer/farther from their foot
          * Reality: they are liking at a pre-recorded clip
      • Results from parametric regression analysis
        • As the spider got closer to the foot, brain activity increased in anxiety-related areas
          • Dorsal anterior cingulate cortex, midbrain nuclei
        • As thee spider got farther from the foot, the brain activity progressively increased in areas that are linked w/ passive coping w/ distant dangers
          • Anterior orbitomedial PFC
      • This reveals the neural circuitry for arachnophobia
    • Davis and Johnsrude 2003
      • Used correlation paradigm to further explain the neural underpinnings of intelligible and unintelligible
      • Intelligibility is on a continuum
        1. Used fMRI, subjects listened to diff stimuli
          * Undistorted condition
          • A: Speech: normal sentences
            * Partially distorted conditions
          • B: Vocoded speech: sentences altered to sound rough but intelligible (similar to Narain et al 2003)
          • C: Segmented speech: sentences altered by dividing the speech stream into separate chunks and replace some parts w/ signal-correlated noise
            • These parts retain many original acoustic features but lack recognizable sounds
          • D: Speech in noise: sentences are placed in the background of continuous speech spectrum noise?
            * Completely distorted condition
          • E: Signal-correlated noise: sentences altered in the same way in the “segmented speech” condition
            • Here, all of the chunks are replaced w/ signal correlated noise -> so the stimuli is completely incomprehensible
              • NOTE: in the 3 partially distorted conditions, there are 3 diff degrees of distortion leading to 3 diff degrees of intelligibility
          • Based on pilot study
          • High intelligibility: 90% of words reported correct
          • Medium intelligibility: 70% of words reported correct
          • Low intelligibility: 20% of words reported correct
        1. The subjects pressed buttons to rate the intelligibility of each stimulus on 4-point scale
      • Results of analyses
        • Subtraction paradigm: compared distorted condition w/ silence
          • Sig bilateral activation in Heschl’s gyrus and surrounding auditory cortices
        • Corelation analyses: see which brain regions respond w/ gradual increase in BOLD signals when the intelligibility of the stimuli increases
        • The following regions show Neural sensitivity to cont variation in intelligibility of stimuli:
            1. superior and middle temporal gyri in LH; extends anteriorly to temporal area and posteriorly angular gyrus
            1. Similar to #1 but distributed more anteriorly in RH
            1. Small part of Broca’s area
      • Follow-up analyses
        • There is a single voxel in the left anterior temporal lobe that was sensitive to intelligibility
          • Activated strongly to undistorted condition
          • Activated weakly in completely distorted condition
          • Activated w/ varying intensities across 3 partially distorted conditions depending on the amount of distortion
            • (i.e. 90%, 70%, or 20% intelligible)
        • IOW: the voxel is not sensitive to the diff ways they are distorted (vocoded, segmented, or embedded in background noise)
    • Take away from this study: brain mechanisms for comprehension of intelligible utterances involve the LH areas mentioned in Narain et al 2003’s study
      • It also involves the RH
    • Here, the correlation paradigm allows rs to discover more brain areas involved
    • Gradual changes in the stimuli taps into diff brain regions
  • Multivariate Pattern Analysis
    • A new paradigm that can decode more fine-grained patterns of brain activity to more precision than the other paradigms
    • In the subtraction and correlation paradigms, each voxel is independent, and the analyses shows separate measures of signal strengths
    • When we use these paradigms to look at the whole brain, it looks at the mean response of each isolated voxel in each condition
    • When we use it to look at the region of interest (ROI), it spatially averages the signals from all the voxels in the region for each condition
    • Limitations: this ignores the relationship b/w voxels, and this can’t detect any info that is latent (dormant)
    • MVPA can identify these dormant patterns, and relate them to cog abilities/tasks
    • Mur et al 2009/ Raizada et al 2010
      • MVPA can help explore whether the perception of similar syllables (ex. ra and la) is associated w/ distinct levels or patterns of brain activity in the superior temporal ROI spanning 9 voxels
      • If the stimuli triggered orthogonal (right angle) 4-voxel activation patterns in ROI, the conventional analysis would miss this
        • Conventional analysis would generate output that shows the same activity for both speech sounds and suggest that ROI cannot discriminate b/w the two
      • MVPA can expose this separate pattern, and suggest the ROI contains at least partially non-overlapping neural voxels that represent the 2 syllables
    • Point: MVPA is very useful
    • Abrams et al 2013
      • Used MVPA to examine the neural substrates that process intelligible vs unintelligible utterances w/ better precision
        1. Present 20 ppl w/ 2 types of stimuli, sequenced in alternating blocks
          * A: Speech (Sp): Intelligible excerpts from famous speeches in 20 C
          * B: Rotated speech (RSp): same experts but spectrally rotated (inverted) as in Narain’s 2003 study
        1. Ppl play close attention to each excerpt and push a button when it ended
      • Rs conducted 2 analyses: Subtraction and MVPA
        • I. Subtraction paradigm/General Linear model (A – B) -> (Sp – RSp)
        • II. MVPA, searchlight version
          • For every voxel in the brain, a 3 x 3 x 3 neighbourhood is made, centred in the given voxel
          • W/in the 27-voxel space, rs determined is they were diff
      • Results
        • Red: conducted subtraction/General linear model (GLM) analysis
          • Areas include 3 left lateral temporal regions (as seen in Narain et al 2003)
          • Most of the bilateral temporal regions (as seen in Davis and Johnsrude 2003)
        • Green: MVPA/ searchlight analysis; Also detected by GLM analysis
          • There are 2 patches where the multi-voxel patterns of activity were sig diff for the two conditions
        • Blue: MVPA/ searchlight analysis; NOT detected by GLM analysis
          • Picked out RH areas the can discriminate b/w the two conditions
          • Located in temporal, parietal, and FL
          • They are associated w/ auditory sensory processing
            • Conclusion: results suggest MVPA can resolve inconsistencies in the literature by being more sensitive in data analysis
31
Q

Electrophysiology

  • 2 Electrophysiological approaches
  • 2 ways to record electrical signals of neurons
  • # 1 Stimulation
    • Penfield 1956
      • How he mapped functional organization of the cortex?
      • Results
        • Stimulate Primary somatosensory cortex
        • Stimulate superior temporal regionstimulated higher-order temporal regions
        • Stimulate language related regions 3 effects
          • Speech arrest
          • Anomia
          • Paraphasia
    • Ojemann 1989 - directly stimulate language-related areas to map out that cortex area
      • Methods - 4 steps
      • 2 main findings
    • 3 cons of e- stimulation
  • # 2 Recording
    • Intracranial
      • fx
      • procedure - 2 steps
      • Creutzfeldt et al 1989 finding
      • Local field potentials
        • What is it
        • 2 ways to measure it
      • Flinker et al 2011 - Studied cortical dynamics of speech perception in 3 epileptic patients
        • Methods
        • Results
    • Extracranial
      • fx
      • EEG
      • ERP
      • 4 ERP parameters
      • Why is +ve or -ve wave deviations don’t matter?
      • Spatial and temporal resolution
      • N400
        • meaning
        • Which electrode can detect N400 w/ greatest amplitude
        • When does it increase
        • Kutas and Hillyard - Showed the waveforms evoked by 3 diff types of sentences presented to subjects on a screen, 1 word at a time
          • 3 conditions
          • results
A

Electrophysiology

  • Both PET and fMRI does not directly measure brain activity; they only track the metabolic consequences
  • 2 Electrophysiological approaches
    • Directly stimulate specific parts in the brain and observe the effects on cognition and b
    • Record electrical signals of neurons
        1. Intracranially: put electrodes in the brain to record the firing of single cells or a cluster of cells
        1. Extracranially: put electrodes n the scalp to record cells firing

Stimulation

  • Penfield 1956
    • Direct electrical stimulation to map the functional organization of the cortex
    • Did this on patients’ w/ epilepsy before removing the brain areas that were thought to cause seizures
    • Used local anaesthesia only as there are no pain receptors in the brain
    • Found that stimulating some areas induce “+ve responses”
      • Ex. stimulate PMC cause involuntary movements in specific body parts
      • Ex. Stimulate Primary somatosensory cortex evoked feelings in specific body parts; they correspond to the somatosensory homunculi
      • Ex. Stimulate occipital regions cause visual hallucination (color, starts, light flickers)
      • Ex. Stimulate superior temporal regions cause auditory hallucinations (ringing, clicking, buzzing)
      • Ex. If he stimulated higher-order temporal regions, this replays the patient’s past experiences that were richer than ordinary recollections
        • flashbacks/dreams
        • This is rare
    • Stimulation to language-related areas can induce “-ve responses”, esp for speech
      • Speech arrest: slowing of speech production
      • Anomia: can’t retrieve a word
      • Paraphasia: distort the semantic or phonological content of a word
        • Ex. say “tephelone” instead of “telephone.”
    • This showed that stimulating specific sites and cause linguistic errors
  • Ojemann 1989
    • directly stimulate language-related areas to study language
    • mapped out the “eloquent” cortex in the left perisylvian territory
    • 120 patients w/ epilepsy
      1. Showed line drawings of familiar objects on a screen in 4-sec intervals
      1. Patient had to name each one using the carrier phrase “this is a ___”
      1. At the beginning of some slides, the rs stimulate one of the several predetermined points on the exposed cortex
        * Each part was stimulated 3 times; w/ all sites were stimulated once b4 any of them were stimulated a 2nd time
      1. Rs provide immediate feedback on patient’s response
        * If stimulating a given site induced error for 2 out 3 trials, that site is essential for naming
    • 2 main findings
        1. For most cases, stimulation disrupted naming as a few discrete sites
          * In 70% patients, 2+ of these areas were identified: 1 in FC, other in temporal/parietal cortex
        1. Rs did stimulation mapping on 90 patients in anterior and posterior perisylvian zones
          * 20% had frontal naming sites
          * 15% had temporal and parietal naming sites
          * Main point: precise naming sites in the brain vary across patients
          * There was only 1 site where errors were evoked more than 50% of the time (located in posterior part of Broca’s area)
  • Some say electrical stimulation is the “gold standard for brain mapping”
  • Cons:
      1. we don’t fully understand the physiological cons of passing current through the cortex
      1. The intensity of stimulation that is needed to identify “eloquent” areas differ across patients and regions
      1. Stimulation to a given site can propagate to other remote brain regions
  • Need to interpret these findings w/ caution!

Recording

  • Intracranial
    • Some rs record neural activity in particular regions
    • It helps doctors to precisely localize the source of epileptic seizures
    • Procedure
      1. Implant electrodes in the tissue that is suspected to surround the epileptogenic site
      1. close the patient’s head and record neural activity for several days cont
        * If the seizure occur, we can identify the point of origin
    • Some rs determine whether and how the firing rates of neurons in the implanted tissue are modulated (regulated)
      • They study the neural activity in single cells of cell assemblies
    • Creutzfeldt et al 1989
      • Found that individual neurons in the right superior temporal gyrus are sensitive to certain features of auditorily perceived words (ex. phoneme, syllables, morphemes)
      • Ex. when the patient listened to a list of multisyllabic words, the firing rate of one cell increased sig in response
      • The critical sounds are velar consonants (i.e. /k/, /g/) combined w/ /r/ or /s/
        • Ex. Christmas, crocodile
      • When the entire list of words was presented again after 2 min, the cell’s response to each word was similar to the response in the 1st time
    • Local field potentials: the sum of the extracellular voltage fluctuations that indicate the sum of events in the dendrites for a local population of neurons
      • This is measured when the patient is doing linguistic tasks
      • Methods
      • Option A: Use electrodes that penetrate many layers of cortex and white matter/ or target subcortical nuclei
      • Option B: use high-density multi-electrode grid, place it directly over the cortical surface to collect data (aka electrocorticography)
    • Flinker et al 2011
      • Studied cortical dynamics of speech perception in 3 epileptic patients
        1. Placed an electrode grid on patient that covers the posterior lateral surface of the left superior temporal gyrus
        1. Ppl listened to 2 types of stimuli
          * I. Syllables (for 150 ms): /ba/, /da/, /ga/
          * II. 3 kinds of words spoken by the same talker
          • 23 pseudowords (3 phonemes)
          • 23 real words (3 phonemes)
          • 4 proper names (5 phonemes)
      • Results:
        • Electrode A (top row): high frequency neural activity response only to words
        • Electrode B (bottom row; 4mm away to electrode A): high frequency neural activity response to both words and phonemes
        • Most activity starts after 200 ms
      • Point: electrocorticography is useful
  • Extracranial
    • Electrical potential produced by large populations of neurons are strong enough to be conducted passively though tissues of the brain, skull, and scalp
    • This is a method to record patterns of activity in a non-invasive manner
    • You place electrodes on the surface of the scalp and compare the voltage fluctuations here w/ a reference location (ex. mastoid bones behind the ears)
    • Electroencephalogram: recording on the changing electrical activity; data is from an electrode on the scalp
      • Has rhythmic undulations (ups and downs) that vary in amplitude, duration; it depends on whether the subject is excited, relaxed drowsy or in a stage of sleep
    • EEG is a collection of many diff neural activity; so it doesn’t provide info on mental operations
      • Rs examine the neural correlates of specific cog capacities
      • Rs examine how the EEG recorded at diff electrodes change when ppl are doing the tasks
    • Event-related potentials (ERPs): EEG patterns that are measured when the stimuli is presented
    • 4 ERP parameters
        1. Latency: time-point, milliseconds after the stimulus; happens when a particular deflection of waveform begins/reach its peak
        1. Amplitude: strength of an effect (mV)
        1. Polarity: whether a deflection is +ve-going or -ve-going
        1. Topography: the scalp distribution of an effect; involved the electrode positions at where the effect was observed
    • Some labs plot -ve up and +ve down OR vv
    • When a wave deviates +vely or -vely, it usually is not sig
      • This is b/c polarity is affected by many irrelevant factors
        • Ex. location of reference electrodes
        • Ex. orientation of the source of scalp recorded signals
        • Ex. if a batch of cortical neurons have their dendrites point to the surface and simultaneously get excitatory input, the resulting electrophysiological activity create current “dipoles”
          • Each dipole has a -ve end near the dendrites and a +ve end near the cell body
        • The summation of these dipoles is detected at the scalp as a -ve voltage
        • This does not mean the operations are done by the neurons, it’s just activity on detected on the scalp
      • Since most ERP studies compare waveforms from the experimental to control condition, the diff in polarity maybe related to cog processes
      • The key thing is there is a polarity diff; the +ve/-ve doesn’t matter
    • Topography
      • Studies vary on the # of electrodes used and their size
      • As the size increases, the difference of ERP effects along the scalp increases
      • The larger range of channels, the easier it is to apply data analysis to somewhat infer the neural underpinnings of ERP effects
      • Inverse problem: ERP has poor spatial resolution but v good temporal resolution
      • Another issue: the electrical activity recorded at the scalp can come from any neural generator
    • N400
      • N = negative
      • 400 = a peak that appears 400 ms post-stimulus
      • centroparietal electrode can detect a N400 w/ the greatest amplitude
      • N400 can track the gradual build-up of semantic content during receptive sentence processing
        • The amplitude increases as we increase the difficulty of integrating the meaning of a word into the preceding context
      • Kutas and Hillyard 1980
        • Showed the waveforms evoked by 3 diff types of sentences presented to subjects on a screen, 1 word at a time
        • Condition 1: all sentences are normal
          • Ex. It was his first day at work
        • Condition 2: all of the sentences ended w/ a word that was semantically anomalous in the given context
          • Ex He spread the warn bread w/ socks
        • Condition 3: all the sentences ended w/ a word that was orthographically incongruent in the given context
          • Ex. She put on her high heeled SHOES
        • Results: final word in anomalous condition trigger a large N400
          • This indicates there was effortful conceptual processing, not surprise
        • The final word in the orthogonal condition is associated w/ a P560
          *
32
Q

Magnetoencephalography

  • Which technique is it related
  • Mechanism
  • ERFs
  • What sensors are used?
  • What does SQUIDS stand
  • 2 Pros of MEG/ERF
  • 2 Cons of MEG/ERF
    *
A

Box 2.2: Magnetoencephalography

  • Non-invasive brain mapping technique related to EEG/ERP
  • Electrical currents associated w/ neural activity create tiny magnetic fields, these fields are recorded at the scalp and time-locked to when the stimuli is presented
  • This produces event-related fields (ERFs)
  • In MEG systems, we detect ERFs w/ 200 sensors called SQUIDS
    • SQUIDS: superconducting quantum interference device
  • Pros of MEG/ERF
    • Outstanding temporal resolution (in ms)
    • Provide good spatial resolution, can help neurosurgeons identify where the seizure happens in epileptic patients
      • This is possible b/c magnetic fields are not distorted as they pass through the brain, skull, and scalp
      • Magnetic field strength dips from their source in a systematic manner
  • Cons
    • It is sensitive to neural activity in the gyri but NOT the sulci
    • Too $$$
      • This is b/c ERFs are tiny signals that are a billionth the size of earth’s magnetic field
      • The lab has to be heavily shielded from outside magnetic fields
      • SQUIDS need to be stored in liquid helium
  • More studies are using this despite the cost
33
Q

Transcranial Magnetic Stimulation

  • 2 reasons why it is not often used
  • What can it do
  • Is it safe?
  • Mechanism
  • Temporal and spatial resolution
  • What doe researchers use as a benchmark
  • 2 strategies are used to determine the appropriate position of a coil (can use 1 or both)
  • Major unkown issue
  • Define online vs offline application of TMS
  • Gough et al - Used TMS and found the complementary roles of 2 diff sectors in the left interior frontal gyrus (LIFG) in making semantic and phonological judgements
    • Methods
    • Results
      *
A

Transcranial Magnetic Stimulation

  • It is very invasive and restricted to those w/ a history of nro dysfunction -> TMS is not used often
  • TMS can quickly alter the organization of neural activity in certain cortical regions by placing magnetic device on the scalp
  • You can facilitate or supress the operation of the target area
  • If safety protocols are followed, TMS is harmless; w/ may cause seizures
  • It can help offset clinical disorders like stroke-induced aphasia, and study language/MD

How It Works

  • Uses EM iniduction
  • Faraday
    • When an electrical current passed thru a wire, it generates a magnetic field that varies w/ time
    • If wire 2 is placed nearby, the magnetic field induces an electric current flow in wire 2
  • For TMS, “wire 1” is the stimulating coil”; “wire 2: is the targeted region of the brain
  • Common coil type: figure-8 shape, current flows around 2 adj parts and sum up at the intersection
  • The flux/magnetic field lines are perpendicular to the plane of the coil
    • When the central part of the coil is placed at a predetermined position on the scalp, the magnetic field that passes through the skull
      • this temp changes the electrophysiological properties of the neurons under the cortical region
    • Temporal resolution: ms
  • Spatial resolution: mm
  • Shifting the coil b/w areas separated by .5 to 1 cm over the PMC can trigger muscle twitches that follow the motor homunculus layout
  • Rs use the intensity threshold for triggering a finger twitch as a benchmark for calibrating the strength of pulses delivered to other regions for high-lv cog processes
  • 2 strategies are used to determine the appropriate position of a coil (can use 1 or both)
      1. Functional localization: move the coil around a general territory until the ability of interest if enhanced/disrupted
      1. Anatomical localization: use neuronavigation system to guide the placement of the coil
        * based on data from structural or fMRI OR a set of standardized coordinates
  • Issues: effects of TMS can spread to remote brain regions; there may be other b cons
    • There are many factors that induce facilitatory or inhibitory effects
  • One main parameter: frequency of pulses
  • Ex. 2 single pulses separated by less than 5 ms -> intracortical inhibition
  • Ex. 2 single pulses separated by 10 – 30 ms -> intracortical facilitation
  • Some pulses can be applied to subjects “online” (during the task performance)
  • Some pulses are applied “offline” (prior to task performance)
  • Ex. Common technique – apply repetitive pulses to target site cont for several min b4 task
    • Reduce excitability of that region after stimulation
    • This can affect subsequent performance
  • TMS cons are measured in RT or accuracy

Applications to Language

  • TMS helps us understand more about language-related brain regions
  • Gough et al 2005
    • Used TMS and found the complementary roles of 2 diff sectors in the left interior frontal gyrus (LIFG) in making semantic and phonological judgements
      • Rostral (pars orbitalis)
      • Caudal (pars opercularis)
    • Subjects did 3 task w/ visually presented pairs of letter strings
      • Synonym judgement – ex. decide if idea and notion hv the same meaning
      • Rhyme judgement – ex. decide if eye and fly end w/ the same sound
      • Visual judgement – ex. decide if txbufr and txbufr have the same characters
    • There are 15 pseudo-randomly organized blocks of 10 trials, w/ the trials in each block involving the same type of task
    • 40% of the trials
      • There are 3 TMS pulses separated by 100 ms and starting 100 ms post-stimulus onset
      • These were delivered to the rostral or caudal of LIFG
    • There are equal # of TMS trials for each task; no more than 2 TMS trials consecutively
    • Results
      • Double dissociation emerged b/w semantic and phonological tasks based on the function of the site of stimulation
      • Judgements on the meanings of word pairs are delayed by rostral LIFG stimulation, not caudal LIFG
      • Judgements on the sound of word pairs were sig delayed by caudal LIFG stimulation, not rostral stimulation
      • Judgements on the appearance of letter strings were not affected by either site
    • There are 2 target areas in the LIFG that make diff contributions to lexical processing
    • These areas are close together; TMS can modify brain activity of brain areas that are close together
      *
34
Q

Major Strengths and Weaknesses of the Different Methods

  • Method
  • Strength
  • Weakness
A
35
Q

What is language

  • Compare and contrast
    • Zebra Finches vs humans
    • Monkey Alarm clocks and Zebra Finches
      *
A

What is language

  • Examples of communication (i.e., there needs to be some interaction between multiple individuals, usually of the same species, to communicate info):
  • Ex. Zebra Finches: male communicating (mate calling) with a female
    • The sounds are sequences of predictable sounds (aka motives)
    • There is enough similarity b/w Zebra finches and humans when communicating
    • It lacks some aspects of communication (ex. we don’t know what is being said)
  • Monkey Alarm Calls: aka notifying each other of a predator
    • Monkeys have different alarm calls for different predators
      • Alarm call 1: leopard alarm call
      • Monkeys from different species recognize each other’s alarm calls
      • They will move closer b/c once the leopard knows it has been spotted, it will give up
      • Unlike the zebra finch, we know what is being said (leopard is here)
      • Communication here is more sophisticated
  • These are not perfect examples of language
  • Main point: there is a continuum of communication; the more complex form is used in humans
36
Q

Neural basis of language

  • What is the brain’s language network made of?
  • What are these other systems also used for?
  • How do we understand words: 5-step
A

Neural basis of language

  • The brain’s language network is formed by a combination of areas (networks) of brain regions that have become differentially specialized for language, but that clearly also relate to other basic cognitive abilities
  • Different systems are used in different aspects of language; these brain areas are also used in other functions (non-language)
    • Vision (ex. reading)
    • Audition (ex. listening)
    • cross-modal associations
    • motor outputs (ex. writing, Brail)
    • planning/executive control (ex. produce a response)
  • How do we understand words?
    1. Extract words from auditory segment
    1. Separate them into phonemes
    1. Associate these to words
    1. Put words together to determine its meaning
    1. Produce a response
  • If certain language functions are present in other species (like birds and monkeys), these functions may evolved to be more specialized in humans
37
Q

A few (of many!) properties of language

  • 6 main properties
A

A few (of many!) properties of language

  • “words” convey “meaning” (lexical semantics)
    • Ex. monkeys has a specific alarm for leopards (basic meaning)
  • Grammar, morphology, and syntax (particular words in particular order)
    • Ex. not present in monkey language (subject, verb, object structure; location of leopard)
  • Generative (i.e., we can create new sentences that everyone can understand)
    • Ex. not seen in monkey speech
  • Specialized for communication
    • Ex. human language is used for communication specifically
    • Ex. Hammer is not specialized for communication – can be used as an alarm, or banging on a pot for music
  • BUT, it stills builds upon other basic abilities like vision, audition, and motor systems, to name a few
  • Other important properties?
38
Q

Cognitive Neuroscience of Language

  • 3 major domains for this discipline
  • What are the 4 other areas
  • How of other areas contribute to this realm?
    • Evolution / Cross-species comparisons
    • Historical analyses (e.g., of language evolution)
      • Study: Compare Spanish in Europe vs South America
        • Looked at hominins (ambiguous words)
        • Define hominins
        • Method
        • Results
    • Computational Modeling
      • Neural network simulation
      • 3 Pros
    • Genetics
A

Cognitive Neuroscience of Language

  • Fundamentally, an interdisciplinary domain, comprised of at least 3 major domains:
    • Cognitive psychology
    • Neuroscience (Neuropsychology = precursor to NRO)
      • Neuropsychology: post-mortem study on what aspects of the patient is impairing language
    • Linguistics
  • Although many other areas make their appearance as well:
    • Evolution / Cross-species comparisons
      • Behavioral, neural levels
      • Some rodents can detect similar sound frequencies as humans
    • Historical analyses (e.g., of language evolution)
      • Tells us…
        • how words change meanings over time
        • how new words are come about
        • Why different communicative abilities evolve
        • what words are central to communication; what words are less relevant
      • Study: Compare Spanish in Europe vs South America
        • Looked at hominins (ambiguous words) – these words have different meaning based on the context they are in
        • Ex. river BANK vs money BANK
        • Results:
          • 50% variability in the language remains the same in two populations
          • 50% variability in the language - the meaning has changed
          • Reason: distinct cultural env, ext env, colonization in South America changed some meanings of words
    • Computational Modeling
      • Neural network simulation: dev an algorithmic or math model of how language works in the brain
      • Pros
        • Develop complex theory
        • Allows us to make predictions for empirical studies
          • Helps us to know what to study next
        • Allow us to do things that may be unethical on real subjects
          • Ex. damage a brain part in the model, and see how the model reacts to the damage
          • Ex. examine if therapy works for brain damage
            • Give different types of therapy to brain damaged model and see how it reacts to each one of it
            • Then test and apply this to humans, and see if it works
    • Genetics
      • Looks at how single and combination of genes leads to different amounts of NT, which in turn may impact our language processing
      • Ex. Diff types of dyslexia may linked to different collection genes; this can be related to whether the guy or gal contributed the genes
39
Q

Classic examples

  • Broca vs Wernicke aphasia
  • Corresponding BA #s?
  • Which hemisphere is more specialized for language?
    • which type of language?
  • Audiovisual integration
    • low lv vs high lv?
  • Amodal semantic combination
  • Articulatory planning
  • Volition
    *
A

Classic examples[EL1]

  • Broca’s and Wernike’s Aphasias
  • In many ways, the seeds of the current interdisciplinary science of language research
  • Behavioural Manifestation:
    • Broca: meaning intact, but not the sentence structure
      • BA. 44 and 45
    • Wernicke: no meaning, fluid speech (word salad)
      • BA 42
  • Very different language impairments, linked to different brain impairments

A basic map of some language systems

  • Note: Many areas serve more than one language relevant or non-relevant ability
  • Most of our cortex influences our language
  • LH is more specialized language, esp visual language
    • Visual inputs, auditory inputs, audiovisual integration
    • Audiovisual integration
      • Low level: how movement of lips correspond to different sounds
      • Higher level: read text on the screen (subtitles) interact with the dialogue the actors is speaking in the movie
    • Amodal semantic combination: how we map info from one modal to another
      • I pat a dog, it feels furry; and you can hear the word “fur”
    • Articulatory planning: plans how to move our mouth to produce specific sounds
    • Volition: decide what we want to produce
  •  Language processing and production is a very complex task, yet our brain has come to do things very efficiently and effectively

[EL1]Understand the difference between the two aphasias

Know where the brain regions are located

40
Q

Tools of the trade

  • Neuropsychology
    • 3 main ways
  • Psycholinguistics/behavior studies
    • McGurk effect
      • Which 2 systems are used?
      • What does this indicate?
A

Tools of the trade

  • Language processing is extremely rapid (ex. when you hear multiple words per second) and takes place across a distributed set of brain regions
  • These brain regions also subserve other abilities (e.g., auditory system is not only for language)
  • What types of methods can be used to probe language?

Neuropsychology

  • We study patients with language impairments, link those to brain damage, usually originally primarily through post-mortem analyses
  • but sometimes we study living individuals like Phineas Gage).
  • Now, the brain is also explored through non-invasive techniques except in special circumstances (e.g., implant electrodes in pre-surgical epileptic patients’ brain).

Behavioural Studies (aka Psycholinguistics)[EL1]

  • E.g., the McGurk effect
    • We use different information from the visual and auditory modality to perceive an ambiguous stimulus
  • Can provide important insights and constraints on cognitive and neural theories of processing
  • These types of tasks are often used in conjunction with other cognitive neuroscience techniques (e.g., ERP).
  • [EL1]Understand how there are different systems contributing to language at the same time, some may be more powerful or relied upon more
  • McGurk effect is a great example of this
41
Q

Tools of the trade cont 2

  • ERP
  • Pro, con
  • Steinhauer, 2014, Applied Linguistics
    • Method
    • Results
A

ERP

  • Behavioural studies are precursors of imaging studies
  • Pros: Excellent for studying fast processing, which is common in language (we need good temporal resolution; helps us track changes over time).
  • Cons: Relatively poor spatial resolution (without convergent evidence from other techniques/studies, data alone only tell you roughly which quadrant of the brain the signal came from).
    • Don’t know the precise location where the signal is coming from

Example from Steinhauer, 2014, Applied Linguistics

  • Method:
      1. Put on EEG cap
      1. Listen to sentences
        * Ex. John ate broccoli at dinner
        * Ex. John ate democracy at dinner
  • Results
    • Blue vertical line: onset/when the words (i.e. broccoli or democracy) are spoken
    • Results: brain signals look similar overall, except at N400 (blue = difference)
    • N400: (400ms after onset)
      • There’s a difference b/w processing the word that makes sense vs one that doesn’t make sense
      • N400 relates to how comprehensible the sentence is
42
Q

Example of Combining Behavioural and Neuroimaging Techniques

  • Baart, Armstrong, Martin, Carreiras, and Frost (2017)
  • Exp 1
    • Methods
  • Exp 2
    • Methods
    • How is it different from Exp 1
    • Purpose
  • 4 noise conditions
  • Results of behavioral exp 1
    • Accuracy & RT
    • Is having a lot noise as bad as predicted, why?
  • Results = ERP Exp 2
    • Analysis
    • What it suggests
  • 4 main insights
A

Example of Combining Behavioural and Neuroimaging Techniques

  • From Baart, Armstrong, Martin, Carreiras, and Frost (2017)
    • Method[EL1]
      • Exp 1: behavioural experiment
          1. See and hear a Spanish word (ex. Apio)
          1. Press a button if what they see and hear match
        • Participants had to go through 4 conditions
          • no noise
          • audio noise
          • visual noise
          • audio and visual noise
      • Exp 2: same experiment but measure subject brain activity w/ EEG
        • Only difference: there is a 1250 ms b/w read + listen to word and pressing the button
        • Purpose: the actual act of pressing the button will activate the motor cortex
        • this delay allows researchers to differentiate b/w when the brain is reading + listening to the word vs pressing the button
  • Noise conditions
    • No noise: white text on black background
    • Visual noise: white dots on the “white text and black background”
        • Audio noise
      • Ex. play a random audio noise -> most ppl don’t know what word it is saying
      • When someone told me the word is “actress”
        • It seems obvious the word is actress in retrospect
      • This suggests that when the audio information is not clear, presenting the word visually can clear up the ambiguity
    • A + V noise (have audio and the visual noise presented)
  • Results – behavioral exp 1
      • Accuracy & RT: Ppl are the most accurate and fastest when there is no noise
    • Auditory noise makes ppl slightly less accurate and a bit slower
    • If you have more noise (i.e. A + V noise), you are the least accurate and the slowest
      • But the effect is not as bad as it seems
        • When we look at RT, AV noise is significantly slower compared to no noise
        • For accuracy, the drop is not as bad
  • Results = ERP Exp 2
    • Researchers did a lot of t-tests
    • After controlling for this (the orange and blue), we get the results (black)
    • The effects happen around N400 (~500 ms)
    • This suggests that the integration of audio and visual noise is happening at the understanding/later semantic level (not early perceptual level)

Integrating insights

  • Combined data point to cross-modal integration occurring at a relatively late time-point, at the level of lexical/semantic processing
    • (combining info from 2 modalities happen later)
  • Surprisingly, the constraints from sublexical units (individual letters / sounds) seems relatively weak. (most integration happens at lexical semantic lv)
  • Potentially due to language tested (Spanish).
    • Transparent orthography: every letter maps onto a particular sound, and vv
    • Results may not apply to other languages that do not have transparent orthography (ex. Eng, Hebrew)
  • Points to value of Cross-linguistic comparisons another type of study that transcends any particular technique
    • *

[EL1] Understand these experiments well and the significance of their results