Neuroling - 1 EGG

NEUROLINGUISTICS
daniele.panizza@gmail.com
NEUROLINGUISTICS
NEUROLINGUISTICS
relevant informa4on about the course

NEUROLINGUISTICS
program is nego4able (I’m at your service)

NEUROLINGUISTICS
provide feedback (I’m at your service)

NEUROLINGUISTICS
slides and readings will be uploaded a"er or before every lecture

NEUROLINGUISTICS
ask ques4ons!
interrupt me any 4me you don’t understand something
NEUROLINGUISTICS
ask ques4ons!
relevant informa4on about me

NEUROLINGUISTICS
ask ques4ons!

psycholinguist (eye-tracking, ERPs, acquisi4on)
NEUROLINGUISTICS
ask ques4ons!

relevant informa4on about you
NEUROLINGUISTICS
ask ques4ons!

relevant informa4on about you
introduce yourself
NEUROLINGUISTICS
papers & textbooks
and maybe others…

what is
NEUROLINGUISTICS
?
what is
NEUROLINGUISTICS
?
Neurolinguistics is the study of the neural mechanisms in
the human brain that control the comprehension, production,
and acquisition of language. As an interdisciplinary field,
neurolinguistics draws methods and theories from fields such
as neuroscience, linguistics, cognitive science, neurobiology,
communication disorders, neuropsychology, and computer
science.
what is NEUROLINGUISTICS ?
Alexander Luria (‘50s)
Harry Whittaker
(Brain & Language, 1974)
two ways to conceive this term
Harry Whittaker
1. neuroscience of language
Harry Whittaker
Neuroscience is the scientific study of the nervous system.
Harry Whittaker
how language is implemented in the central nervous system
Harry Whittaker
perception = hearing/reading
Harry Whittaker
production = speaking (Brain & Language, 1974)
understanding = semantics, concepts

Harry Whittaker
2. NEURO - LINGUISTICS
Harry Whittaker
phonology/morphology
syntax
(formal) semantics
pragmatics
Harry Whittaker
phonology/morphology
syntax
(formal) semantics
pragmatics
we will do a li-le bit of both
overview of the course 13 classes + final exam
tenta1ve program
lessons 1-2 lesson 5-6-7
SYNTAX in the brain (fMRI)
warming-up & intro
SEMANTICS and CONCEPTS (fMRI)
history, morphology of the brain
PRAGMATICS (fMRI)
physiology of the nervous system
methods in neuroscience lessons 8
NEUROLINGUISTICS electrified
lessons 3-4 (EEG and MEG studies)
neuroscience of language syntax (P600) and seman4cs (N400)
parsing and on-line sentence processing
hearing, reading, wri4ng the N400 debate
neuroimaging lessons 9-10
phonology
formal neuroseman4cs (nega4on,
composi4on, modals, nega4ve
polarity items)
pragma4cs & implicatures
what is the brain?
what is the brain?
what is the brain?
Aristotle (350 bc)

what is the brain?
Aristotle (350 bc)
the ra4o of brain size to body size was greatest in more

intellectually advanced species, such as humans
what is the brain?
Aristotle (350 bc)

cogni4on as product of…

what is the brain?
Aristotle (350 bc)

cogni4on as product of… HEART!

what is the brain?
Aristotle (350 bc)

BRAIN as cooling system

(the smarter you are, the more cooling you need)
what is the brain?
Aristotle (350 bc)


Galen (150 ad)

what is the brain?
Aristotle (350 bc)


Galen (150 ad) observa4on of lesions in injured roman gladiators

what is the brain?
Aristotle (350 bc)



nerves project to and from the brain
what is the brain?
Aristotle (350 bc)



mental experiences reside in the…

what is the brain?
Aristotle (350 bc)



mental experiences reside in the… VENTRICLES!

Vesalius (1500)
Anterior
how the brain looks..

commissure
Midbrain
Cerebellum
Brainstem Pons
Medulla Spinal cord
(E) (F)
Corpus Corpus Thalamus Basal ganglia
Internal
Cerebral cortex callosum capsule Lateral callosum
ventricle Caudate
(gray matter) Putamen
White Globus
matter Internal pallidus
Caudate capsule
Third
ventricle
Putamen
Tail of
caudate
nucleus
Temporal Lateral
lobe ventricle
(temporal
Optic
horn) Mammillary
Anterior chiasm
commissure Basal forebrain body
Amygdala nuclei Hippocampus Fornix
Connec&ng the Hemispheres:

Grey vs White Ma.er Corpus Callosum
brain: reference system
Superior (above)
Anterior Posterior
(in front of) (behind)
Inferior (below)
Left Right
Lateral Medial Lateral
Posterior
(behind)
which one is… dorsal? ventral? caudal? rostral? anterior? posterior?


dorsal
dorsal
ventral
dorsal
caudal
ventral
dorsal
rostral caudal
ventral
dorsal
rostral caudal
anterior
ventral
dorsal
posterior
rostral caudal
anterior
ventral
brain: cortex
brain: cortex
cortex is grey (grey mager)

brain: cortex

it contains cell bodies (astrocytes & neurons)
brain: cortex

it represents the computa4onal power of the brain (lots of computers)
brain: cortex

highly convoluted folded sheet

brain: cortex


why?
brain: cortex


why? because for evolu4onal purposes we need do stuff
more neurons into a non-expanding skull
(the benefits are manifold)
brain: cortex


gyrus/giri = bumps
brain: cortex


gyrus/giri = bumps
sulcus/sulci = grooves
cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure
Temporal lobe Cerebellum

cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure

cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure

cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure

cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure

cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure

cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure

cortex morphology
Central Parietal
sulcus lobe
Frontal Occipital
lobe lobe
Sylvian
fissure

what do you know about the brain?
what is the length of an adult’s brain?
14 cm 8 cm
24 cm 20 cm
what is the length of an adult’s brain?
14 cm 8 cm
24 cm 20 cm
how many neurons are there in a brain?
100 billion 455 million
20 billion 84 billion
how many neurons are there in a brain?
100 billion 455 million
20 billion 84 billion
which animal has the biggest brain?
whale elephant
dolphin man
it’s not the size that matters!
some brains..
which animal has the higher brain-to-body ratio ?
whale ant
dolphin man
hgp://io9.gizmodo.com/5890414/the-4-biggest-myths-about-the-human-brain
To circumvent the limitations of a basic, brain-to-body mass ratio, researchers devised a more complex measurement, known as the
"encephalization quotient" (or "EQ"), that measures the ratio of an animal's brain and body size relative to other, similarly sized animals. In
doing so, EQ not only takes into account that brain size tends to increase with body size, but that brain size does not necessarily increase at
the same rate as body size.
neuron
26 CHAPTER 2 • NEURONS AND GLIA
The Golgi Stain

The Nissl stain, however, does not tell the whole story. A Nissl-stained neu-
ron looks like little more than a lump of protoplasm containing a nucleus.
Neurons are much more than that, but how much more was not recog-
nized until the publication of the work of Italian histologist Camillo Golgi
(Figure 2.2). In 1873, Golgi discovered that by soaking brain tissue in a sil-
ver chromate solution, now called the Golgi stain, a small percentage of
neurons became darkly colored in their entirety (Figure 2.3). This revealed
that the neuronal cell body, the region of the neuron around the nucleus
that is shown with the Nissl stain, is actually only a small fraction of the
total structure of the neuron. Notice in Figures 2.1 and 2.3 how different
histological stains can provide strikingly different views of the same tissue.
Today, neurohistology remains an active field in neuroscience, along with
its credo: “The gain in brain is mainly in the stain.”
The Golgi stain shows that neurons have at least two distinguishable
parts: a central region that contains the cell nucleus, and numerous thin
tubes that radiate away from the central region. The swollen region con-
FIGURE 2.2 taining the cell nucleus has several names that are used interchangeably:
Camillo Golgi (1843–1926). cell body, soma (plural: somata), and perikaryon (plural: perikarya). The
(Source: Finger, 1994, Fig. 3.22.) thin tubes that radiate away from the soma are called neurites and are of
two types: axons and dendrites (Figure 2.4).
The cell body usually gives rise to a single axon. The axon is of uniform
diameter throughout its length, and if it branches, the branches generally
extend at right angles. Because axons can travel over great distances in the
body (a meter or more), it was immediately recognized by the histologists
of the day that axons must act like “wires” that carry the output of the
neurons. Dendrites, on the other hand, rarely extend more than 2 mm in
soaked brain 4ssue in a silver chromate solu4on: neurons in their beauty Soma
diameter throughout its length, and
extend at right angles. Because axons
neuron body (a meter or more), it was imme

of the day that axons must act like
neurons. Dendrites, on the other han
26 CHAPTER 2 • NEURONS AND GLIA
The Golgi Stain

The Nissl stain, however, does not tell the whole story. A Nissl-stained neu-
ron looks like little more than a lump of protoplasm containing a nucleus.
Neurons are much more than that, but how much more was not recog-
nized until the publication of the work of Italian histologist Camillo Golgi
(Figure 2.2). In 1873, Golgi discovered that by soaking brain tissue in a sil-
ver chromate solution, now called the Golgi stain, a small percentage of
neurons became darkly colored in their entirety (Figure 2.3). This revealed
that the neuronal cell body, the region of the neuron around the nucleus
that is shown with the Nissl stain, is actually only a small fraction of the
total structure of the neuron. Notice in Figures 2.1 and 2.3 how different
histological stains can provide strikingly different views of the same tissue.
Today, neurohistology remains an active field in neuroscience, along with
its credo: “The gain in brain is mainly in the stain.”
The Golgi stain shows that neurons have at least two distinguishable
parts: a central region that contains the cell nucleus, and numerous thin
tubes that radiate away from the central region. The swollen region con-
FIGURE 2.2 taining the cell nucleus has several names that are used interchangeably:
Camillo Golgi (1843–1926). cell body, soma (plural: somata), and perikaryon (plural: perikarya). The
(Source: Finger, 1994, Fig. 3.22.) thin tubes that radiate away from the soma are called neurites and are of
two types: axons and dendrites (Figure 2.4).
The cell body usually gives rise to a single axon. The axon is of uniform
FIGURE 2.3
Golgi-stained neurons. (Source: Hubel, 1988, p. 126.)
soaked brain 4ssue in a silver chromate solu4on: neurons in their beauty Soma
neuron
neurons are cells
Soma
Dendrites Neurites
Axon
: Hubel, 1988, p. 126.)
FIGURE 2.4
The basic parts of a neuron.
neuron
neurons are cells
func4ons: process informa4on

Soma
Dendrites Neurites
Axon
: Hubel, 1988, p. 126.)
FIGURE 2.4
neuron
neurons are cells

Soma
transmit informa4on
Dendrites Neurites
Axon
: Hubel, 1988, p. 126.)
FIGURE 2.4
neuron
neurons are cells

Soma
transmit informa4on
body = soma
Dendrites Neurites
Axon
: Hubel, 1988, p. 126.)
FIGURE 2.4
neuron
neurons are cells

Soma
transmit informa4on
body = soma
Dendrites Neurites
neurites Axon
: Hubel, 1988, p. 126.)
FIGURE 2.4
neuron
neurons are cells

Soma
transmit informa4on
body = soma
Dendrites Neurites
neurites Axon
denrites (several): INPUT
: Hubel, 1988, p. 126.)
FIGURE 2.4
neuron
neurons are cells

Soma
transmit informa4on
body = soma
Dendrites Neurites
neurites Axon

axon (one): OUTPUT
: Hubel, 1988, p. 126.)
FIGURE 2.4
neuron
neurons are cells

Soma
transmit informa4on
body = soma
Dendrites Neurites
neurites Axon

axon (one): OUTPUT
(white mager
contains axons!)
: Hubel, 1988, p. 126.)
FIGURE 2.4
bipolar, and if there are three or more, the cell is multipolar. Most neurons
in the brain are multipolar.
neuron
Classification Based on Dendrites
Dendritic trees can vary widely from one type of neuron to another. Some
have inspired elegant names like “double bouquet cells.” Others have less
interesting names, such as “alpha cells.” Classification is often unique to a
types of neurons
particular part of the brain. For example, in the cerebral cortex (the struc-
ture that lies just under the surface of the cerebrum), there are two broad
classes: stellate cells (star-shaped) and pyramidal cells (pyramid-shaped)
(Figure 2.20).
Soma
Unipolar
Multipolar
Bipolar
FIGURE 2.19
Classification of neurons based on the number of neurites.
bipolar, and if there are three or more, the cell is multipolar. Most neurons
in the brain are multipolar.
neuron
Classification Based on Dendrites 46 CHAPTER 2 • NEURONS AND GLIA
Dendritic trees can vary widely from one type of neuron to another. Some
have inspired elegant names like “double bouquet cells.” Others have less Another simple
dendrites have spin
interesting names, such as “alpha cells.” Classification is often unique to a not are called aspin
types of neurons
particular part of the brain. For example, in the cerebral cortex (the struc- For example, in th
cells, on the other
ture that lies just under the surface of the cerebrum), there are two broad
classes: stellate cells (star-shaped) and pyramidal cells (pyramid-shaped) Classification B
(Figure 2.20). Information is deliv
rites in the sensory
the eye. Cells with t
Other neurons hav
Soma mand movements;
the nervous system
to this classification
Stellate cell
Classification B
Some neurons hav
Unipolar the other; these are
neurons have short
body; these are cal
cerebral cortex, for
extend to other pa
In contrast, stellate
cortex and are ther
Classification B
The classification sc
Multipolar neurons as reveale
scientists to identify
resulted in a schem
example, the motor
the neurotransmitte
also classified as ch
Bipolar transmitter. Collect
up the brain’s neur
FIGURE 2.19
Classification of neurons based on the number of neurites. ▼ GLIA
We have devoted
While this decision
roscientists conside
day, they suppose,
tantly to informatio
At present, howev
function mainly by
be subordinate, wit
Pyramidal cell
neuron: synapse
INTRODUCING THE BRAI
A neuron KEY TERM

Neuron
A type of cell tha
up the nervous s
and supports, am
other things, cog
function.
Cell body
Part of the neuro
containing the n
and other organe
Dendrites
Branching struct
carry information
other neurons.
Neurons consist of three basic features: a cell body, dendrites that receive information and Axon
neuron: synapse
1. electrical current propagates down the axon

INTRODUCING THE BRAI
A neuron KEY TERM

Neuron
A type of cell tha
up the nervous s
and supports, am
other things, cog
function.
Cell body
Part of the neuro
containing the n
and other organe
Dendrites
Branching struct
carry information
other neurons.
Neurons consist of three basic features: a cell body, dendrites that receive information and Axon
1. Microtubules do not extend into the terminal.
neuron: synapse 2. The terminal contains numerous small bubbles of membrane, called
synaptic vesicles, that measure about 50 nm in diameter.
3. The inside surface of the membrane that faces the synapse has a partic-
ularly dense covering of proteins.
4. It has numerous mitochondria, indicating a high energy demand.
1. electrical current
propagates down the axon
Presynaptic
axon terminal
Mitochondria
reaches the synap;c bu=on Synapse
Synaptic FIG
vesicles Th
te
or
im
na
Postsynaptic dendrite fro
Synaptic Ne
cleft re
Receptors of
syn
1. Microtubules do not extend into the terminal.
neuron: synapse 2. The terminal contains numerous small bubbles of membrane, called
synaptic vesicles, that measure about 50 nm in diameter.
3. The inside surface of the membrane that faces the synapse has a partic-
ularly dense covering of proteins.
4. It has numerous mitochondria, indicating a high energy demand.
1. electrical current
propagates down the axon
Presynaptic
axon terminal
Mitochondria
reaches the synap;c bu=on Synapse
small space between

Synaptic FIG
vesicles Th
te
AXON or
im
na
Postsynaptic dendrite
DENDRITES / SOMA Synaptic
fro
Ne
cleft re
Receptors of
syn
neuron: synapse
end of an axon beginning of a dendrite

(presynap1c) (postsynap1c)
neuron: synapse

2. ac4on poten4al
reaches the axon terminal
neuron: synapse

2. ac4on poten4al
3. chemicals are
released into the
synap4c clem
(neurotransmigers)
neuron: synapse

2. ac4on poten4al
3. chemicals are
released into the
synap4c clem
(neurotransmigers)
4. neurotransmigers
bind to the receptors
in the postsynap4c
dentrites or soma
neuron: synapse
1.Neurotransmitters
3. Electrical
2. Electrical
neuron: synapse
5. synap4c poten4al is conducted passively (without ac4on

poten4al)
1.Neurotransmitters
3. Electrical
2. Electrical
neuron: synapse

poten4al)
6. if passive currents are sufficiently strong they cause an ac4on

poten4al in the neuron
1.Neurotransmitters
3. Electrical
2. Electrical
neuron: synapse

poten4al)
6. if passive currents are sufficiently strong they cause an ac4on

poten4al in the neuron
7. dendrite poten4als sum up (passive current short ranged)
1.Neurotransmitters
3. Electrical
2. Electrical
time, thus facilitating a burst of hormone secretion into the circulation. The nels connect to one another, creating
fact that gap junction pores are large enough to allow molecules such as ATP electrical continuity between the two
neuron: synapse
and second messengers to diffuse intercellularly also permits electrical syn- cells. (B) Rapid transmission of signals
apses to coordinate the intracellular signaling and metabolism of coupled at an electrical synapse in the crayfish.
An action potential in the presynaptic
cells. This property may be particularly important for glial cells, which form
neuron causes the postsynaptic neuron
large intracellular signaling networks via their gap junctions. to be depolarized within a fraction of a
millisecond. (B after Furshpan and Pot-
ter, 1959.)
(A) Presynaptic (B)

cell membrane
25 Presynaptic
Connexons neuron
Membrane potential (mV)

−25
−50
Postsynaptic 25
cell membrane 3.5 nm Postsynaptic
20 nm neuron
0
Pores connecting
cytoplasm of two −25
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
pre-synap4c poten4al
(A) Presynaptic (B)
cell membrane
25 Presynaptic
Connexons neuron

−25
−50
Postsynaptic 25
20 nm neuron
0
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
(A) Presynaptic (B)
cell membrane
AXON Connexons 25 Presynaptic
neuron

−25
−50
Postsynaptic 25
20 nm neuron
0
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
(A) Presynaptic (B)
cell membrane
AXON 0 or 1 Connexons 25 Presynaptic
neuron

−25
−50
Postsynaptic 25
20 nm neuron
0
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
(A) Presynaptic (B)
cell membrane
neuron
0
propagates as electric current along the axon

−25
−50
Postsynaptic 25
20 nm neuron
0
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
(A) Presynaptic (B)
cell membrane
neuron
0

−25
ACTIVE (regenerates) −50
Postsynaptic 25
20 nm neuron
0
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
(A) Presynaptic (B)
cell membrane
neuron
0

−25
ACTIVE (regenerates) exploits myelin (electric isola4on) −50
Postsynaptic 25
20 nm neuron
0
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
(A) Presynaptic (B)
cell membrane
neuron
0

−25
reaches dendrites or somas

Postsynaptic 25
20 nm neuron
0
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)
pre-synap4c poten4al (A) Presynaptic (B)

cell membrane
neuron
0

−25

triggers the release of Postsynaptic
neurotransmigers (few
cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
eccitatory or inhibitory) 0 neuron
Pores connecting
neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting
long-ranged (e.g. from motor area to spine chord)

neurons
−50
Brief (~0.1 ms)
synaptic delay
0 1 2 3 4
Time (ms)
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting

neurons
−50
Brief (~0.1 ms)
synaptic delay
post-synap4c poten4al 0 1 2
Time (ms)
3 4
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting

neurons
−50
Brief (~0.1 ms)
synaptic delay
Time (ms)
3 4
DENDRITES or SOMA
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting

neurons
−50
Brief (~0.1 ms)
synaptic delay
Time (ms)
3 4
DENDRITES or SOMA
influenced by excitatory or inhibitory forces, PASSIVE (does not regenerate)
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting

neurons
−50
Brief (~0.1 ms)
synaptic delay
Time (ms)
3 4
DENDRITES or SOMA
it is gradable/con4nuous
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting

neurons
−50
Brief (~0.1 ms)
synaptic delay
Time (ms)
3 4
DENDRITES or SOMA
it is gradable/con4nuous short-ranged
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting

neurons
−50
Brief (~0.1 ms)
synaptic delay
Time (ms)
3 4
DENDRITES or SOMA
benefits of the sum of all surrounding poten4als
neuron: synapse
ter, 1959.)

cell membrane
neuron
0

−25

cell membrane
types,
3.5 nm
25
Postsynaptic
20 nm
Pores connecting

neurons
−50
Brief (~0.1 ms)
synaptic delay
Time (ms)
3 4
DENDRITES or SOMA
benefits of the sum of all surrounding poten4als
is triggered by ion-channels bound to receptors (many types)
neuron: action potential
.qrk(75-100).ps 11/30/05 1:23 PM Page 83
▼ THE ACTION POTENTIAL, IN REALITY
neuronal membrane at res1ng state electric poten1al INTRODUCIN
g >> g
K Na
Outside
Sodium Potassium in
cell
K+ channel K+ channel
+ + + + + + + + + + + + + + + + + + + V
m
o
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
(a) The action potential consists of a number of phases.

each neuron is surrounded by a cell membrane that acts as a barrier to

the passage of chemicals (Na+ = sodium, K+ = potassium)
.qrk(75-100).ps 11/30/05 1:23 PM Page 83
g >> g
K Na
Outside
Sodium Potassium in
cell
+ + + + + + + + + + + + + + + + + + + V
m
o
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell


res4ng poten4al = -70mV (inside nega4ve, outside posi4ve)

.qrk(75-100).ps 11/30/05 1:23 PM Page 83
g >> g
K Na
Outside
Sodium Potassium in
cell
+ + + + + + + + + + + + + + + + + + + V
m
o
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell


res4ng poten4al = -70mV (inside nega4ve, outside posi4ve)

.qrk(75-100).ps 11/30/05 1:23 PM Page 83
ion channels (= voltage gates) are found only in axons:

generate ac4on poten4al!
g >> g
K Na
Outside
Sodium Potassium in
cell
+ + + + + + + + + + + + + + + + + + + V
m
o
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell

g >> g
K Na
Outside
Sodium Potassium
cell
+ + + + + + + + + + + + + + + + + + + V
m
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
(a) depolariza1on (posi1ve current in) electric poten1al INTRODUCIN
g >> g
Na K
in
K+ K+
V
m o
– –
Na+ Na+
Sodium influx – 80 mV
The action potential consists of a number of phases.
(b)
if passive current is sufficiently strong:

g >> g
K Na
Outside
Sodium Potassium
cell
+ + + + + + + + + + + + + + + + + + + V
m
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
g >> g
Na K
in
K+ K+
V
m o
– –
Na+ Na+
(b)

g >> g
K Na
Outside Na+ channelsSodium
open (posi4ve electric
Potassium flux inside the cell)
cell
+ + + + + + + + + + + + + + + + + + + V
m
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
g >> g
Na K
in
K+ K+
V
m o
– –
Na+ Na+
(b)

g >> g
K Na
cell
voltage reaches -50mV
+ + + + + + + + + + + + + + + + + + + V
m
– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
g >> g
Na K
in
K+ K+
V
m o
– –
Na+ Na+
(b)

g >> g
K Na
cell
voltage reaches -50mV
+ + + + + + + + + + + + + + + + + + + V
m
at this point the membrane becomes completely permeable:
CHARGE
– – – – – – – INSIDE
– – –ABOUT
– – –TO– CHANGE
– – – –RAPIDLY
–
(ac4on Kpoten4al)
+ K+
Inside – 80 mV
cell
g >> g
Na K
in
K+ K+
V
m o
– –
Na+ Na+
(b)
V
m

– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
(a)
g >> g
Na K
K+ K+
V
m
– –
Na+ Na+
(b) posi1ve charge inside the cell (maximum spike) electric poten1al INTRODUCIN
g >> g
K Na
Potassium efflux in
K+ K+
– – – – – – –
V
m
o
+ + + + + + +
– 80 mV
(c) The action potential consists of a number of phases.

V
m

– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
(a)
voltage goes beyond 0 mV (posi4ve charge)
g >> g
Na K
K+ K+
V
m
– –
Na+ Na+
g >> g
K Na
Potassium efflux in
K+ K+
– – – – – – –
V
m
o
+ + + + + + +
– 80 mV

V
m

– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
(a)
g >> g
Na K
ACTION
K+ POTENTIAL! K+
V
m
– –
Na+ Na+
g >> g
K Na
Potassium efflux in
K+ K+
– – – – – – –
V
m
o
+ + + + + + +
– 80 mV

V
m

– – – – – – – – – – – – – – – – – – –
Inside
K+ K+
– 80 mV
cell
(a)
g >> g
Na K
ACTION
K+ POTENTIAL! K+
V
charge inside the cell is reversed (from neg to pos)m
– –
Na+ Na+
g >> g
K Na
Potassium efflux in
K+ K+
– – – – – – –
V
m
o
+ + + + + + +
– 80 mV

V
m

– –
Na+ Na+
(b)
g >> g
K Na
Potassium efflux
K+ K+
– – – – – – –
V
m
+ + + + + + +
ion pumps restore the gradients of K+ and Na+ (burning

– 80 mV glucose through ATP)
(c) polariza1on of the cell (coming back to res1ng state) electric poten1al INTRODUCIN
g >> g
K Na
K+ K+
in
+ + + + + + + + + + + + + + + + + +
V
m o
– – – – – – – – – – – – – – – – – –
K+ K+
– 80 mV
(d) Time
V
m

– –
Na+ Na+
K+ ion channels become open
(b)
g >> g
K Na
Potassium efflux
K+ K+
– – – – – – –
V
m
+ + + + + + +

g >> g
K Na
K+ K+
in
+ + + + + + + + + + + + + + + + + +
V
m o
– – – – – – – – – – – – – – – – – –
K+ K+
– 80 mV
(d) Time
V
m

– –
Na+ Na+
(b) nega4ve poten4al of the cell is restored via outward flow of K+

g >> g
K Na
Potassium efflux
K+ K+
– – – – – – –
V
m
+ + + + + + +

g >> g
K Na
K+ K+
in
+ + + + + + + + + + + + + + + + + +
V
m o
– – – – – – – – – – – – – – – – – –
K+ K+
– 80 mV
(d) Time
V
m

– –
Na+ Na+

g >> g
charge inside the cell is reversed again (from pos to K Na
Potassium efflux
neg) K+ K+
– – – – – – –
V
m
+ + + + + + +

g >> g
K Na
K+ K+
in
+ + + + + + + + + + + + + + + + + +
V
m o
– – – – – – – – – – – – – – – – – –
K+ K+
– 80 mV
(d) Time
V
m

– –
Na+ Na+

g >> g
Potassium efflux
neg) K+ K+
– brief–hyperpolariza4on,
– – undershoot
– –(more nega4ve
– than at rest)
V
m
+ + + + + + +

g >> g
K Na
K+ K+
in
+ + + + + + + + + + + + + + + + + +
V
m o
– – – – – – – – – – – – – – – – – –
K+ K+
– 80 mV
(d) Time
V
m

– –
Na+ Na+

g >> g
Potassium efflux
neg) K+ K+
– brief–hyperpolariza4on,
– – undershoot
– –(more nega4ve
– than at rest)
V
m
prevents the ac4on poten4al from traveling back

+ (cannot
+ depolarize
+ again
+ straight
+ away,
+ refractory
+ period)
g >> g
K Na
K+ K+
in
+ + + + + + + + + + + + + + + + + +
V
m o
– – – – – – – – – – – – – – – – – –
K+ K+
– 80 mV
(d) Time
2. The substance must be released in exogenous transmitter mimics the post- neurotransmitters.

response to presynaptic depolarization, and
the release must be Ca2+-dependent.
Demonstrating the identity of a neurotransmitter at a synapse requires showing (1) its pres-
ence, (2) its release, and (3) the postsynaptic presence of specific receptors.
(1) (2) (3)
Action
potential
1 Neuro-
transmitter Presynaptic
present terminal
Application of
transmitter, agonists,
or antagonists
Ca2+
Ca2+
2 Neurotransmitter 3 Neurotransmitter
Postsynaptic cell
released receptors activated

(1) (2) (3)
Action
potential
1 Neuro-
present terminal
Application of
or antagonists
Ca2+
Ca2+
Postsynaptic cell
transmigers are short-lived


(1) (2) (3)
Action
potential
1 Neuro-
present terminal
Application of
or antagonists
Ca2+
Ca2+
Postsynaptic cell

they are destroyed and recycled (reuptake)

(1) (2) (3)
Action
potential
1 Neuro-
present terminal
Application of
or antagonists
Ca2+
Ca2+
Postsynaptic cell

they are destroyed and recycled (reuptake)
otherwise, they would over-excite neuron (cf. cocaine)
neurotoxins
neurotoxins
fasciculins
blocks the Ach transmiger
in the receptor (cannot be broken
down by acetylchilinesterase (AChE)
(involuntary muscle contrac4on)
toxins are enough to inhibit a cholinergic synapse. Botu- phates. These are irreversible inhibitors of AChE, and by

linum toxins are extraordinarily specific enzymes that de- preventing the degradation of ACh, they probably kill their
stroy certain of the SNARE proteins in the presynaptic victims by causing a desensitization of ACh receptors.
terminals, which are critical for transmitter release (see The organophosphates used today as insecticides, like
Box 5.3). Ironically, this specific action of the toxins made parathion, are toxic to humans only in high doses.
them important tools in the early research on SNAREs.
Although its mechanism of action is different, black
widow spider venom also exerts deadly effects by affect-
neurotoxins ing transmitter release (Figure A). The venom first in-
creases, and then eliminates, ACh release at the neuro-
fasciculins muscular junction. Electron microscopic examination of
synapses poisoned with black widow spider venom re-
veals that the axon terminals are swollen and the synap-
tic vesicles are missing. The action of the venom, a pro-
tein molecule, is not entirely understood. Venom binds
with proteins on the outside of the presynaptic mem-
brane, perhaps forming a membrane pore that depolarizes
the terminal and allows Ca2! to enter and trigger rapid
and total depletion of transmitter. In some cases, the
venom can induce transmitter release even without the
need for Ca2!.
The bite of the Taiwanese cobra also results in the
blockade of neuromuscular transmission in its victim, but
by yet another mechanism. One of the active compounds FIGURE A
in the snake’s venom, called "-bungarotoxin, is a peptide Black widow spiders. (Source: Matthews, 1995, p. 174.)

need for Ca2!.
tetrodotoxins dentrotoxins
blocks the ion exchange


need for Ca2!.

no polariza4on/depolariza4on

need for Ca2!.

no polariza4on/depolariza4on
the amplitude of ac4on poten4al does not vary

what varies is the frequency of the spikes (ac4on

poten4al per second = spiking rate)

spiking rate = informa4onal code (010101)


where does the informa4on processing take place?



1. at the postsynaptic space: between
where does the informa4on processing take place? transmitters and receptors


receptors are transmitter-gated ion

channels (flow of K+, Na+. Cl-)


receptors are transmitter-gated ion

channels (flow of K+, Na+. Cl-)
they create a synaptic potential


2. inside the neurons: they sum up the excitatory receptors are transmitter-gated ion
and inhibitory information from various inputs channels (flow of K+, Na+. Cl-)
(coming from action potentials) they create a synaptic potential


excitatory neurotransmigers (ACh, DA): make the neuron more likely to fire as it is more
posi4ve than normal


excitatory neurotransmigers (ACh, DA): make the neuron more likely to fire as it is more
BEARc05.qrk(101-132).ps 11/30/05 1:27 PM Page 128
posi4ve than normal

inhibitory neurotransmigers (e.g. GABA): make the neuron less likely to fire as it is more nega4ve
than normal128 CHAPTER 5 • SYNAPTIC TRANSMISSION
Excitatory synapse Inhibitory synapse

(active) (inactive)
Dendrite
Soma
Axon hillock
Record Vm Record Vm
let’s take a brake
What do neurons use to talk to each other?

let’s take a brake
What do neurons use to talk to each other?
A cellular phone.
let’s take a brake hgp://neuroscien4stryangosling.tumblr.com
let’s take a brake hgp://neuroscien4stryangosling.tumblr.com
back to history…
1800
Phrenology
Gall (1800)
Phrenology
Gall (1800)
two assump4ons:
Phrenology
Gall (1800)
two assump4ons:
1. different regions of the brain perform different func4ons
and are associated with different behaviours
Phrenology
Gall (1800)
two assump4ons:
1. different regions of the brain perform different func4ons
and are associated with different behaviours
2. size of those regions produces distor4ons of the skulls

and correlates with individual differences in cogni4on and personality
Holism VS. Phrenology
Flourens (1830)
Flourens (1830)
experimental abla4on in birds

Flourens (1830)

provided experimental evidence that
cerebellum and brain have different func4ons
Flourens (1830)

But:
Flourens (1830)

But:
1. the skull is not correlated with

the shape of the brain
Flourens (1830)

But:
1. the skull is not correlated with

the shape of the brain
2. all the regions of the brain par4cipate

equally in all cerebral func4ons
(holis4c view)
What about all those bumps on the brain’s surface? Do they perform
Functional specialization: Broca ferent functions as well? The idea that they do was irresistible to a y
Austrian medical student named Franz Joseph Gall. Believing that bu
on the surface of the skull reflect the bumps on the surface of the brain
proposed in 1809 that the propensity for certain personality traits, su
generosity, secretiveness, and destructiveness, could be related to th
Broca (1860, but see Dax) mensions of the head (Figure 1.10). To support his claim, Gall and hi
lowers collected and carefully measured the skulls of hundreds of pe
representing the extensive range of personality types, from the very g
to the criminally insane. This new “science” of correlating the structu
the head with personality traits was called phrenology. Although the c
of the phrenologists were never taken seriously by the mainstream scie
community, they did capture the popular imagination of the time. In fa
textbook on phrenology published in 1827 sold over 100,000 copies.
One of the most vociferous critics of phrenology was Flourens, the
man who had shown experimentally that the cerebellum and cereb
perform different functions. His grounds for criticism were sound. For
thing, the shape of the skull is not correlated with the shape of the b
In addition, Flourens performed experimental ablations showing that
ticular traits are not isolated to the portions of the cerebrum specifie
phrenology. Flourens also maintained, however, that all regions of the
brum participate equally in all cerebral functions, a conclusion that
was shown to be erroneous.
The person usually credited with tilting the scales of scientific opi
firmly toward localization of function in the cerebrum was French ne
ogist Paul Broca (Figure 1.11). Broca was presented with a patient
FIGURE 1.11 could understand language but could not speak. Following the man’s d
Paul Broca (1824–1880). By carefully in 1861, Broca carefully examined his brain and found a lesion in th
studying the brain of a man who had lost frontal lobe (Figure 1.12). Based on this case and several others lik
the faculty of speech after a brain lesion (see Broca concluded that this region of the human cerebrum was specif
Figure 1.12), Broca became convinced that dif-
ferent functions could be localized to different
responsible for the production of speech.
parts of the cerebrum. (Source: Clarke and Solid experimental support for cerebral localization in animals qu
twosecretiveness,
generosity, pa4ents that could not speak
and destructiveness, could be related to th
lowers collected and carefully measured the skulls of hundreds of pe
twosecretiveness,
lowers one hadandright
collected hemiplegia
carefully measured the skulls of hundreds of pe
twosecretiveness,
but other func4ons were intact
the head(including language
with personality understanding)
traits was called phrenology. Although the c
twosecretiveness,
with personality understanding)
traits was called phrenology. Although the c
the they
community, casedidofcapture
Tan (he could imagination
the popular say only ‘Tan’)
of the time. In fa
twosecretiveness,
with personality traits wasunderstanding)
called phrenology. Although the c
the they
community, casedidofcapture
Tan (he could imagination
the popular say only ‘Tan’)
of the time. In fa
conclusion:
One of the most this regioncritics
vociferous is responsible for Flourens,
of phrenology was speaking the
ge 11 on the surface of the skull reflect the bumps on the surface of the brain
two
generosity, pa4ents
secretiveness, that could not
and destructiveness, speak
could be related to th
the
community, case
they
▼ THE didof TanOF(he
capture
ORIGINS the could imagination
popular
NEUROSCIENCE
say only11‘Tan’)
of the time. In fa
conclusion:
currents to a circumscribed region of perform different functions. Hissulcus
Central grounds for criticism were sound. For
dog could elicit discrete movements.
peated these experiments with mon- ticular traits are not isolated to the portions of the cerebrum specifie
al of this same region of the cerebrum phrenology. Flourens also maintained, however, that all regions of the
larly, German physiologist Hermann brum participate equally in all cerebral functions, a conclusion that
presented evidence that the occipital The person usually credited with tilting the scales of scientific opi
required for vision. firmly toward localization of function in the cerebrum was French ne
ok, we now know that there is a very ogist Paul Broca (Figure 1.11). Broca was presented with a patient
um,FIGURE
with1.11different parts performing could understand language but could not speak. Following the man’s d
ps studying
of thethefunctional divisions
brain of a man who had lost of the frontal lobe (Figure 1.12). Based on this case and several others lik
te of
the those
faculty of produced by the
speech after a brain lesion phrenol-
(see Broca concluded that this region of the human cerebrum was specif
ke ferent
the functions
phrenologists, scientists
could be localized to differenttoday
ge 11 on the surface of the skull reflect the bumps on the surface of the brain
two
generosity, pa4ents
secretiveness, that could not
and destructiveness, speak
could be related to th
the
community, case
they
▼ THE didof TanOF(he
capture
ORIGINS the could imagination
popular
NEUROSCIENCE
say only11‘Tan’)
of the time. In fa
conclusion:
currents to a circumscribed region of perform different functions. Hissulcus
Central grounds for criticism were sound. For
dog could elicit discrete movements.
peated these experiments with mon- ticular traits are not isolated to the portions of the cerebrum specifie
al of this same region of the cerebrum phrenology. Flourens also maintained, however, that all regions of the
larly, German physiologist Hermann brum participate equally in all cerebral functions, a conclusion that
presented evidence that the occipital The person usually credited with tilting the scales of scientific opi
required for vision. firmly toward localization of function in the cerebrum was French ne
ok, we now know that there is a very ogist Paul Broca (Figure 1.11). Broca was presented with a patient
um,FIGURE
with1.11different parts performing could understand language but could not speak. Following the man’s d
ps studying
of thethefunctional divisions
brain of a man who had lost of the frontal lobe (Figure 1.12). Based on this case and several others lik
te of
the those
faculty of produced by the
speech after a brain lesion phrenol-
(see Broca concluded that this region of the human cerebrum was specif
ke ferent
the functions
phrenologists, scientists
could be localized to differenttoday
Functional
4.1 Processes: thespecialization: Broca
neuroanatomy of language
History and eary locationist theories
Paul Broca Leborgne („Tan“) Lelong

1824 – 1880 neuro syphilis stroke
•  apparently preserved •  apparently preserved

comprehension comprehension
•  only articulations: „tan“, •  production limited to five

„tan-tan“ words (‘oui’, ‘non’, ‘toi’,
‘toujours’ and ‘Lelo’)
•  died eight days after
admission •  died one year post-stroke
•  post-mortem inspection of •  post-mortem inspection of

the brain the brain
Structures Formelles du Langage 03.02.2016

Functional specialization: Broca
4.1 Processes: the neuroanatomy of language

History and eary locationist theories
M. Lelong!
M. Leborgne !

Inferior Frontal Gyrus
Broca’s Areathe
4.1 Processes: (BA 44, 45)
neuroanatomy (pars
of opercularis, aka frontal operculus,
language
Broca‘s Area and pars triangularis)

Functional specialization: Wernicke
Wernicke (1874)
Wernicke (1874) pa4ent that could speak very fluently

but he said things that made no sense

he did not understand anything

he did not understand anything
Broca s area: Wenicke s area:

Production of Comprehension
Speech of Speech
classical model (Wernicke-Geschwind)
Broca’s Aphasia: speech impaired but good comprehension

Wernicke’s Aphasia: comprehension impaired but fluent speech

ncorporating this new evidence does not necessarily entail the abandon-
idea of anclassical
amodal store ofmodel (Wernicke-Geschwind)
semantic concepts. Mahon and Caramazza
ntain that an amodal system is at the core of semantic memory but that

EENTH-CENTURY MODELS OF SPEECH
Wernicke’s Aphasia: comprehension impaired but fluent speech
providing
cific
in the
around
renology
ost the
terances
ca
a
e patient
The Lichtheim model of speech and aphasia links together
the
Classical Model
Broca‘s Aphasia
fluentness, prosody,
Conduction Aphasia
articulation, word finding, and both comprehension and production preserved BUT
complex grammar disturbes repetition impaired
comprehension more or less
preserved
Wernicke’s Aphasia
comprehension disturbed
prosody preserved, production undisturbed to
overshooting


Criticisms to classical Model
•  Functional-anatomical classification can not (or not sufficiently) account for clinical observations
•  Broca’s aphasia can be found with no lesion to Broca’s area
•  Isolated cortical lesions of the inferior frontal gyrus often lead to transient symptoms, not
corresponding to Broca’s aphasia
•  Productive speech deficits also after lesions to Wernicke’s area
•  Very rarely comprehension deficits after unilateral damage to Wernicke’s area

a spoiler for the next classes
Modern hypothesis on Broca‘s Area
phonological
processing
syntactic
processing
semantic
processing

back to the brain..
3.4 A short primer into the Human Brain
Gross Anatomy

more on Functional specialization: Brodmann
Brodmann (1909)
Brodmann (1909)
Cytoarchitectural map of cortex
Brodmann (1909)
each area with a common architecture

was given a number
Brodmann (1909)
each area with a common architecture

was given a number
why do brain areas have different

cytoarchitectonic proper4es?
why do brain areas have different cytoarchitectonic proper4es?

Cor$cal Layers (Cytoarchitecture)
I Dendrites of deeper cells
II Small granule cells
III Variety of cells, many pyramidal in shape
IV Mainly granule cells

V Pyramidally shaped cells
larger than in layer III
VI Heterogeneous layer of neurons

blends into white matter
White Matter
more on functional specialization: Penfield
Penfield (1950)
Penfield (1950)
Penfield electrically s4mulated

the cortex (no pain receptors)
Penfield (1950)

electrical s4mula4on of 400

living and awake pa4ents
Penfield (1950)


they were going under

brain surgery for epilepsy
Penfield (1950)



occipital lobe: “a star came down toward my nose”

Penfield (1950)



central sulcus: “those fingers and my thumb gave a jump”

Penfield (1950)



central sulcus: “those fingers and my thumb gave a jump”
temporal lobe: “I hear music again”

similar and roughly resemble a trapeze artist hanging upside down, his
more on functional specialization: somatotopy

legs hooked over the top of the postcentral gyrus and dangling into the
medial cortex between the hemispheres, and his head at the opposite,
lower end of the gyrus (Figure 12.18). A somatotopic map is sometimes
called a homunculus (from the Latin diminutive of “man”; the little man in
the brain).
somatotopic mapabout
Two things are obvious of the body surface
a somatotopic map. First, onto
the mapprimary
is not somatosensory cortex (S1)
always continuous but can be broken up. Notice in Figure 12.18 that the
Trunk
Head
Neck
Elbo
For
Arm
Hip
ear
Leg
Ha
m
Fin
nd
ge
rs
Th
um
b
Ey
e
No
se Foot
Fa
ce
Up Toes
p er
lip
Lips Genitals
Lower li
p
Teeth
Gums
Jaw
gue
Ton
ry nx
ha
al
P
in
om
bd
-a
tra
In
similar and roughly resemble a trapeze artist hanging upside down, his
more on functional specialization: somatotopy

legs hooked over the top of the postcentral gyrus and dangling into the
medial cortex between the hemispheres, and his head at the opposite,
lower end of the gyrus (Figure 12.18). A somatotopic map is sometimes
called a homunculus (from the Latin diminutive of “man”; the little man in
the brain).
somatotopic mapabout
Two things are obvious of the body surface
a somatotopic map. First, onto
the mapprimary
is not somatosensory cortex (S1)
always continuous but can be broken up. Notice in Figure 12.18 that the
Trunk
Head
Neck
Elbo
For
Arm
Hip
ear
w
the homunculus!
Leg
Ha
m
Fin
nd
ge
rs
Th
um
b
Ey
e
No
se Foot
Fa
ce
Up Toes
p er
lip
Lips Genitals
Lower li
p
Teeth
Gums
Jaw
gue
Ton
ry nx
ha
al
P
in
om
bd
-a
tra
In
more on functional specialization of the brain
map of cortical areas
map of cortical areas
to know more..
and if you want to have more fun with brain func4ons..
http://www.fmriconsulting.com/brodmann/Introduction.html
readings:
Ward: chapters 1 and 2
Bear: chapters 1 to 7
credits (to serious Neurolinguists):
Einat Shetreet
Andrea San; Emiliano Zaccarella

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Neuroling - 1 EGG

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NEUROLINGUISTICS

relevant informa4on about the course

relevant informa4on about the course

program is nego4able (I’m at your service)

relevant informa4on about the course

program is nego4able (I’m at your service)

provide feedback (I’m at your service)

relevant informa4on about the course

program is nego4able (I’m at your service)

provide feedback (I’m at your service)

slides and readings will be uploaded a"er or before every lecture

relevant informa4on about the course

program is nego4able (I’m at your service)

provide feedback (I’m at your service)

slides and readings will be uploaded a"er or before every lecture

relevant informa4on about the course

program is nego4able (I’m at your service)

provide feedback (I’m at your service)

slides and readings will be uploaded a"er or before every lecture

relevant informa4on about me

relevant informa4on about the course

program is nego4able (I’m at your service)

provide feedback (I’m at your service)

slides and readings will be uploaded a"er or before every lecture

relevant informa4on about me

relevant informa4on about the course

program is nego4able (I’m at your service)

provide feedback (I’m at your service)

slides and readings will be uploaded a"er or before every lecture

relevant informa4on about me

relevant informa4on about the course

program is nego4able (I’m at your service)

provide feedback (I’m at your service)

slides and readings will be uploaded a"er or before every lecture

relevant informa4on about me

papers & textbooks

and maybe others…

Alexander Luria (‘50s)

Alexander Luria (‘50s)

Alexander Luria (‘50s)

how language is implemented in the central nervous system

how language is implemented in the central nervous system

understanding = semantics, concepts

how language is implemented in the central nervous system

understanding = semantics, concepts

how language is implemented in the central nervous system

understanding = semantics, concepts

how language is implemented in the central nervous system

understanding = semantics, concepts

Aristotle (350 bc)

Aristotle (350 bc)

the ra4o of brain size to body size was greatest in more

Aristotle (350 bc)

the ra4o of brain size to body size was greatest in more

cogni4on as product of…

Aristotle (350 bc)

the ra4o of brain size to body size was greatest in more

cogni4on as product of… HEART!

Aristotle (350 bc)

the ra4o of brain size to body size was greatest in more

cogni4on as product of… HEART!

BRAIN as cooling system

Aristotle (350 bc)