UNIT 5: SOCIAL BEHAVIOR

Introduction

1. Overview

These aren’t the only components of social behavior, there’s also bonding (affection) and oral judgement.

Perception and recognition of social information

1. Overview

Social Perception is the ability to make sense out of people and people interactions from sensory data.

- Visual
- Touch
- Auditory
- Chemical

Other animals like dogs, get a lot of information through licking and smelling thanks to the pheromones, so
that they can know if the other dog is healthy, is showing signs of aggression, if it’s sexualy availabe…
humans don’t have this.

2. Vision –Face processing

Faces convey a great deal of social information.

- Emotional value
- Normatively, we can know the gender of someone
- We can also know where the person's attention is placed on
- We can guess their interests
- Age
- Arousal (the guy on a picture looking at a girl’s butt)
- Outraged (this guy’s girlfriend who caught him)
- By interpreting faces, we can even guess what the situation is

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 - Maybe this guy has done this before and that’s why his girlfriend looks so mad at him. He’s
being disloyal.

Therefore, faces are a really important entry of information about social aspects that will help us understand
others and guide our behavior.

They also guide our first impressions (studies about how we behave with people we’ve just met).

The face and corporal posture are the main elements for a first impression.

We make judgement on whether we can or cannot trust the person

based solely on facial attributes and expressions:

- Distance between the eyebrows

- Rise of the eyebrows
- Lips
- Closure of the eyes

He showed us a video in which they changed these parameters on a guy’s face. It was interesting to see that
even though it was a continuous / gradual morph, we perceived it as a sudden change on the expression /
facial structure. This sudden perception happens when internal systems activate, like the amygdala.

Our judgement on people based on attributes is extremely fast.

- Experiment: They showed 5 different parameters on faces and showed them to people. They had to
say if they found the attractive, likable, competent, trustworthy, etc. Then, gave them free time to
look at the pictures and then asked him how confident they are on their judgement.
- With 100ms we form a first impression in many aspects that will last long. The longer we look,
the more confident we are about our impression.
- If you give a lot of time and more information about the person, then there’s more variability
(respect the first impression). But without the information, there isn’t.

It probably goes through the low route of the amygdala processing (from the Thalamus directly to the
amygdala).

How do we perceive faces?

- By eye tracking
- It measures the direction and fixation of the eyes.
- When a normal person looks at the picture of a face, there are hot spots →
- Eyes
- Mouth
- Nose
- Shape of the hair
- Contorn of the face

The areas that we use to process these objects are located in the INFERIOR TEMPORAL (IT) (the ventral
pathway of the vision) → “what” pathway.

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 - Experiment on a macaque
- IT
- We have neurons that are very selective to faces
- In the brain of a macaque
- Histogram plot of a spiking rate
- How many action potentials the neuron is firing during the presentation
of a stimulus.
- Some neurons will respond very specifically to faces
- Don’t respond to another body part

Study:

There’s a case in which some patients need to have electrodes on the brain (epileptic patients in which
drugs don’t have an effect). The only way to be able to stop suffering epilectic seizures is removing a part
of the cortex, which could be the source of the epileptic activity. Most of the time, this source is located in
the Temporal Lobe, since it’s a highly active region (it has the hippocampus). The thing is that, after removing
it, the patients would have agnosia, hyersexualation, trouble in memory function, etc.

Nowadays they put a net of electrodes on top of the cortex but below the duramater (electrocorticography).
This way, when the patient has a seizure, they know the exact source so that they don’t affect functions as
much. The only way in which an experimenter can have a human with electrodes implanted on a human’s
brain.

In the fusiform gyrus, there are locations in which the neurons react to faces and just a little bit to leaves
(plants) and even less to cars and houses. Other regions may react mostly to faces but also to other parts of
stimuli. Depends on the region.

- Video of a patient, that when his brain is stimulated he sees the doctor’s face morph into someone
else's face.
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 ● FFA (The Fusiform Face Area)
This region of the IT cortex processes face information. Electrical stimulation evokes changes in face
perception
➢ Neurons that like complex stimuli
Experiment (Reward system)
While first-time mothers show activity at the FFA to the sight of the face of their own infant and others, they
show a selective increase in the activity of reward regions (nucleus accumbens) to happy>neutral>sad faces
of their own infants

Present 3 faces (happy, neutral and sad) of babies (their own and other people babies) to first time mothers.
When a person sees a face the mother gets activation in the inferior temporal cortex. When they get faces
that are emotional and we have a contrast between happy faces, neutral and sad faces, and compare the
mother’s own baby against another baby, the nucleus accumbens (pleasure) get more active (mostly neutral
faces, not sad faces) to their own babies, but not to another baby.

Relates the faces to the recognition of who the person is (experiment)

In vision for social information the face is the most complex stimuli that we can extract (a lot of relevant
information)
Reward system - in first time mothers there's activation when they see a face (as it's natural), but what's
important, is that the nucleus accumbens gets more active to their own babies when seeing expressions.
Mostly to happy and neutral faces.

● Amygdala.
The lateral nucleus of the amygdala receives info from the inferotemporal (IT) cortex (visual). This pathway
of recognition of faces flows directly to the amygdala. Amygdala contains several nuclei. Amygdala receives
from many places.
➢ Most are reward or memory (all the things that we think are emotional processes and memory)
➢ Lateral nucleus mostly receives from the sensory thalamus, inferotemporal cortex and sends to
entorhinal cortex (memory and functional inhibition)
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 ➢ Amygdala sends to the striatum (reward), to the hippocampal formation
➢ Central nucleus sends to…
○ hypothalamus (physiological
responses),
○ pons and medulla (basic
common satisfactions)
○ substantia nigra (reward system)
Important - the network of the regions, this is
what determines functions, not the regions
themselves.
➔ Regions in the brain it's always based on
who they get information from, and who
they send information to.

Experiment with monkeys (amygdala):

Neurons will be sensitive to visual information.Scientifics put electrodes in the amygdala of some monkeys.
They record the activity from single neurons. They show them the image of a monkey.

They do histograms and raster plots for action potentials - neuron spike rates.

A) Monkey fixates on a face

B) Neurons with selective responses (activation or inhibition) to parts of the face

They know if the monkey is looking at the ear, etc.

C) Neurons selective to the eyes

They see that the neurons react to eyes and inhibit in other parts - neurons are selective to certain parts of
the face. Doesn't matter if it's a video or a static image.

D) A neuron selective for the face of a particular individual monkey no matter the expression

There's also a neuron in the amygdala of this particular monkey that reacts to a certain face of a specific
monkey.

E) A neuron selective for threat, no matter the monkey

There's another neuron that doesn't care about the owner of the face and is fixated about thread expression
of any monkey.

Regulates attentional direction of the gaze to detect features that convey emotions, specially fear

Patient with SM pattern→ Patient had a lateral amygdala damage. She had the inability to identify other's
expressions, and didn't recognize emotional expressions (fear mostly).
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 - c) In the image there’s no amygdala (a la ressonància)
- b) SM pattern vs. Normal pattern of regular person

When she was told to look into eyes, she regained the ability. Without the amygdala you don't extract the
information unless you are told to.

The amygdala needs a specific pattern to recognise the face.

Interim Summary

1. Faces convey a great deal of social information, being the main element guiding our first impressions →
Trait judgements are established on the first 100ms of seeing a face!

2. Face perception involves the Fusiform Face Area (FFA), at the Inferior Temporal (IT) cortex, which belongs
to the ventral visual pathway, involved in object recognition (the “What” pathway). → Electrical stimulation
of the FFA evokes changes in face perception.

3. The amygdala receives information from faces via a connection with the IT

- It contains some neurons responding to parts of faces; others selective for the face of a particular
individual; others selective to a face expression regardless of the individual.
- Amygdalar lesions disturb attentional guidance of the gaze for facial expression recognition
(especially fear).

4. Reward regions (nucleus accumbens) selectively increase their activity to the sight of happy faces of a
first-time mother’s own infant.

3. Touch –Social pleasant stroking

Many animals have some social behavior containing a lot of physical contact. It’s an affiliative
behavior. Social touch promotes socio-affective wellbeing. We like being touch, affiliative touch.
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● Social touch - many mammals like being touched because:
○ Promotes wellbeing from a social perspective
■ Reduces stress
● Less blood pressure
● Less heart rate
● Less cortisol
■ Reduces pain
● Less substance P
○ One of the mediators substance that go to the bones to receptors, it
releases the information of pain.
■ Calms
● More serotonin
■ Promotes bonding (friendship, parents and children, etc.)
● More oxytocin

➔ Gentle stroking (caress) has a particular via for transmission information (via specific CT afferents)
◆ The CT one (non myelinated and slow).
◆ Some C fibers carry low threshold touch (CT = C Touch fibers)
● These fibers bring gentle touch.
○ Gentle touch (caresses)→ type of touch that is characterised by being...
◆ not very strong nor light
◆ Nor fast nor slow
○ The right combination between velocity and force they fire more, and the
more they fire, more pleasure you get.
○ Slowly moving low force touch (caress) stimulates CT afferents CT afferent
firing rates correlate with perceived pleasantness

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 ■ The axons don't get to somatosensory regions like other kinds of touch, they project
in the insula in a topographic way. CT afferents stimulation is processed at the insula
● Those who lack AB afferent, they don't know where you're touching (also don't
show activity in the somatosensory areas) but they do feel pleasure. If they lack
the CT aference, what they lose is the pleasure.

Interim Summary

1. Social Touch promotes socio-affective well-being → It calms, reduces stress and pain and promotes
bonding.

2. Caresses, a form of social touch, are transmitted via CT afferents → CT afferents respond maximally to
moving touch stimuli within a narrow force range (gentle) and a narrow velocity range (generally slow), and
their activity directly correlates with perceived pleasantness.

3. CT afferent information is processed at the insular cortex

- People lacking CT afferents show activity to caresses at the somatosensory cortex (information
carried by Aβ touch afferents) but not the insula
- People lacking Aβ afferents show activity to caresses at the insula but not the somatosensory cortex
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 - Newborns show insular activity to caresses
- Neurons at the insula are highly selective to moving touch (stroking) velocity.

4. Therefore, caresses seem to be processed by a different domain of touch, characterized not by its sensory-
discriminative functions, but by its social context and accompanying subjective component, with a dedicated
peripheral and central processing system.

SOCIAL COGNITION
1. Introduction

● Social cognition - a set of processes that help us adapt to our social environment. Extremely complex,
there are so many variables.
○ There seems to be a relationship between the brain size of a primate and the number of
members in their group.
■ In other species it correlates with mating systems that favor their bonding.
● The Social Brain Hypothesis
● The evolution of the brain is linked to societal life. In primates, brain size
increases as a function of group size. The larger the group, the higher the
complexity of social relations and behaviors to process
● We are the ones with the largest neocortex and we can keep a lot of
information on there, this is why it is so huge.
● This theory is supported for primates.
○ We don't use the same processes to understand people and objects.
■ Living organisms are not as predictable.
Social information is more complex, but
we try all the time to predict. The brain
is a predictive/inference machine.
○ Ability to understand, store and use efficient
information about people (also ourselves),
interpersonal behavior abilities (what to do),
rules to act adequately.
○ 2 types of mental inferences
■ Short lasting things - transient state
● Who does what, why does he do
it (maybe he drinks to ease his
thirst)?
● Agency, goals and intentions
■ Long lasting characteristics
● Personality traits and social scripts (I know him, so I know he's drinking not
because he's thirsty but because of the social situation/because he's nervous)

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Mirror mechanism
1. Mirror neurons
- Neurons that discharge both when an individual performs a given behaviour and when an individual
observes another person performing the same or a similar behaviour; these neurons are found in
many brain cortical areas of monkeys and other species, including humans, marmosets and bird

● Mirror mechanism - transient state

○ How do we understand others (behavior/emotions)?
■ Humor is about breaking the expectations
● The facial expression of fear of Macacs is similar to our facial expression of
happiness, but these emotions aren’t the same.
○ Ramachandran - perceiving that you still have the arm through a mirror reflecting the other
one, reduces the pain.
○ Rizzolatti - discovered the mirror neurons. They were recording neurons in the motor cortex
that not only respond to a specific action made by the individual, but also to the same action
done by another individual.
○ Iacoboni - Experiments with monkeys. The cell fires when the monkey grabs food and also
when the experimenter is the one grabbing something (even though the monkey is still). The
animal behaves normally”. These discoveries were made when they were studying...
■ Ventral premotor cortex
■ Fire to goal-directed hand movements
■ Also to observing those movements

2. Mirror neuron system (macaque hand-grasping actions)

Are in areas painted in red.

● Superior Temporal Sulcus (STS) and Inferior Temporal

(IT): high-order visual areas involved in motion processing.
Active during observation, not action (no mirror neurons).
Visual processing.
● Parieto-frontal Network (PFG, AIP, F5): receives
information from the high-order visual areas. Contains
neurons with mirror properties. (Mirror neurons (fronto-parietal network) → Lower parietal area →
Lower intraparietal area → Brocca area (F5) → Primary prefrontal sensory area.)
● Primary Motor Area (F1): receives information from the parieto-frontal network. Contains neurons
with mirror properties arranged topographically according to each body part (that means that are
mirror neurons organized).
○ Fronto-parietal network, sensory and motor functions (they active themselves when we see
and execute the action) → Primary Motor Area is needed to do/execute the action.
○ Primary areas → High level areas → Fronto-parietal → Sensory and motor → Primary motor
area → Limb.
○ Some neurons in F5 and F1 suppress their activity during action observation. WHY? Probably,
to prevent our own movements during action observation

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 ● Pre-supplementary motor area (F6) and ventral prefrontal cortex (vPFC): control the parieto-frontal
network.
● Supplementary Motor Area (SMA) and hippocampus contain mirror neurons (single-neuron study
with 21 patients). Some showed facilitation and others suppression to observed actions.

What do neuron mirrors actually respond to?

● Mirror neurons respond both to execution of an action and to the observation of that action, which
helps imitation.
● Keep the goal, change the action. Testing what information mirrors neurons encode:
○ Response of hand-grasping mirror neuron during the execution and observation of hand and
reverse pliers grasping.
○ Response of one hand-gaspring to eat (preferred) and grasping to place (non-preferred)
○ Response of a mirror neuron to the event of a peanut breaking and to grasping a ring.
Regardless of sensory modality, this neuron prefers the peanut breaking event.
● Mirror neurons encode action goals!
● IMPORTANT: many parieto-frontal neurons encode action goals. But mirror neurons also encode
them when observing.
○ The mirror mechanism does nothing but extend the same functional goal-related
organization from action execution to action observation.
● Mirror neurons aid understanding action. Ex: “I hear and I forget, I see and I remember, I do, and I
understand”.
○ Evidence for the role of mirror neurons in action understanding:
■ Action interferes with action execution. Observing incongruent biological motion
inferes with own movement.
■ Motor expertise aids identifying other’s action goals. Premotor cortex of ballet
dancers is more activated when observing ballet than capoeira, and vice versa.
■ Understanding worsens after transient inhibition of the premotor homunculus.
Inhibition of the hand with TMS worsens the identification of hand actions but not lip
actions, And vice versa.
● Mirror neurons aid action understanding! But not all behaviors consist on such obvious actions..

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Affective Empathy

Empathy: ability to put ourselves in the other’s places, to tune to his/her emotions, feeling what the other
feels. To empathize requires to experience the same emotional state, or an isomorphic one, that the one
the person observed (or imagined) is feeling.

Empathy vs. Sympathy

Put yourself in the other person’s shoes vs. Put

yourself in the other person's shoes from
experience.

Empathy vs. Emotional contagion

There are no clear boundaries about the other

person’s feelings and ours.

Cognitive empathy vs. Affective empathy

Comprehension (abstract reasoning) vs. Feeling.

To initiate a social behavior is necessary to have

self constancy.

Lesions at the VMPFC (Ventro Medial Prefrontal Cortex) affect cognitive empathy, but not affective. And
lesions at the IFG (Inferior Frontal Gyrus) affect affective empathy but not cognitive.

Observing emotions (sensory processing) would elicit somatic sensations (visceromotor processes) in the
observers that are similar to those that would occur if the observers themselves were experiencing that
emotion. This could allow the observers to identify others’ emotion by capitalizing on their own sensations
concerning that emotion.

Case example: DISGUST

Insula: sensorimotor, cognitive, chemical senses and social-emotional inputs of information.

● Also, integrates chemical senses with sensorimotor aspects of ingestion (ex. texture of food drives
chewing).
● Retching crates disgust emotions.

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 ● And promotes orofacial affiliative gestures.

Anterior insula is active both to feeling disgust and to observing disgust.

Case example: PAIN

Pain empathy is mediated by the affective qualities of pain, not by its sensory ones.

● Brain regions specific to receiving pain (NOT observing). Sensorimotor cortex, the second
somatosensory area and the causal portion of ACC.
● Brain regions shared between receiving AND observing pain: anterior insula, rostral ACC (both
correlate with empathy ratings) cerebellum and brainstem.

A putative general mirror mechanism would aid emotion understanding. → “Observing an emotion such as
disgust would elicit visceromotor processes and representations in the observers’ brain that are similar to
those processes and representations that would occur if the observers themselves were experiencing that
emotion. This could allow the observers to identify others’ emotion by capitalizing on their own visceromotor
processes and representations concerning that emotion.”

The key nodes of the networks involved seem to be the Anterior Insula (AI) and the Anterior Cingulate
Cortex (ACC).

Cognitive empathy

Ability to understand that other people have beliefs, desires and intentions that not necessarily agree with
ours; to infer and represent the content of these mental processes and to develop theories that enable us
to predict the behavior of a person in front of different environmental situations.

It implies using cognitive functions such as mental flexibility and theory of mind (ToM; mentalizing).

● Theory of mind
○ Hyperactivation of the Temporo-Parietal Junction (TPJ)
■ Inferring the intentions of perceived social movements or social behavior (transient
states).

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 ■ Distinguish social intentions from my own social intentions. Are they the same?
■ Monitors the decentralization process that enables alternating attention between own
mental states and other’s.
○ Hyperactivation of the Medial Prefrontal Cortex (MPFC)
■ It connects to limbic areas. Functions:
● Positive or negative evaluations of self and others (emotional
value: amygdala).
● Distinguishes between one’s intention/beliefs and the beliefs
and intentions of others.
● Hippocampus→ memory retrieval of self and others.
● Active in strategy games when believing to play against a real
opponent (not a computer). Ex: rock, paper, scissors.
● Involved in abstraction, reflexive thinking and in creating temporal realities,
physically or mentally different.
● It realtes to the representation of more stable characteristics related to
personality traits and the application of rules in social scripts.
● It is important in inferring permanent psychological and social characteristics
of people as well as in evaluating and correctly applying social scripts to each
situation.
○ EX: típico ejemplo de que la una niña deja una muñeca en una caja, otra niña la cambia de
caja. Cuando la niña vuelve mira en la caja donde ella la había dejado realmente, no donde
está ahora. Si un niño adivina este hecho, significa que cumple con la teoría de la mente, de
poder ponerse en el estado mental de la otra persona.
○ The ToM network (particularly the MPFC) is independent from the abstract cognitive
reasoning work. Except when abstract reasoning involves a high degree of moral judgement.
○ TOJ and MPFC are active both when inferring the thoughts of others and when reflecting
upon own thoughts.

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 UNIT 6: MEMORY AND LEARNING
1. Introduction
1.1. The persistence of memory

We don’t store memories or experiences in a specific place, rather, the circuits we use to execute an action,
perceive a stimulus, etc. change. It consolidates changes in the structure of the nervous system, supporting
the networks enabling us to perceive, act, think and plan.

Every time we activate a network to do something, depending on the degree of activity of the network, it
changes (a lot or a bit) everytime we activate it. The concept of learning is that these processes we use to
do things are subject to change by experience.

- Slightly different perception of what we’ve already seen

- Emit different responses
- Changing the way we process information
- We’re adapting ourselves

The change in circuitry that we use to process things is called learning.

When the change becomes lasting, it’s what we call memory. That's why memories are so distributed in our
cortex. Remembering would be replaying the circuits (every time we replay it, it can change as well, that’s
why it can change a lot with time).

1.2. Learning → memory

When we process things, there’re structural and functional changes → Synaptic plasticity

- Kids have more of this plasticity

- When it’s “permanent” we call it memory

1.3. Hebbian theory

The more the circuit is active, the stronger the connection within the circuit gets.

“Let us assume that the persistence or repetition of a reverberatory activity (or "trace") tends to induce
lasting cellular changes that add to its stability. ... When an axon of cell A is near enough to excite a cell B

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and repeatedly or persistently takes part in firing it, some growth process
or metabolic change takes place in one or both cells such that A's
efficiency, as one of the cells firing B, is increased.”

- Hebb, D.O. (1949).The organization of behavior.

Years later, Eric Candel found the first experimental evidence.

Hebb → “Neurons that fire together, wire together” (not exactly true,
one should lead a little bit in time → spike-timing-dependent
plasticity)

- When a neuron (A) takes part in firing another one (B), and B is
already being fired by another neuron (C), the connexion
(between A andB) is going to be stronger.

1.4. Call assemblies and dual-footprint hypothesis

The brain does not work by individual synapses, but by cell assemblies, which
are big neuronal groups being active simultaneously through recurrent
connections.

Cell assemblies lie at the basis of neural representation and through plastic
changes they support learning and memory (engrams). 1 neuron can
participate in many different engrams.

Therefore, memories appear distributed in the brain across different neural

networks. Specific stimuli can activate elements within the network (not
necessarily all) that, through recurrent connectivity, ignite the engram
granting access to the complete neural representation of that memory.

Dual-footprint hypothesis: Memory formation (engrams) is a two-step

process.

- Step 1: Experience generates a brief reverberating activation pattern

in neural networks (cell assemblies)
- Step 2: If the activation reaches a threshold, a long-term memory
footprint can be formed (permanent change).

1.5. Classification of memory types

When we talk about processes of learning and

memory, we can make a quantitative qualification
according to time.

**To know → everything related to semantic

memory is not linked to the context, at some point
in your life you learn these.

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Explicit knowledge is that one that you could actually explain with words, while the implicit you can’t. Riding
a bike for example, for example, can’t be learnt by an explanation, there’s no point in explaining. Instead,
you can say that you’ve just learnt about the classification of the types of memory and tell someone.

Explicit memory needs less trials yet it is subject to external influences (for example after an exam, when we
forget what we’ve studied or if the teacher told us that this classification is wrong and that we should forget
it).

The limbic system (hippocampus, fornix, etc.) takes part in explicit memory.

2. Implicit learning and memory (more stable)

Implicit learning and memory is independent of consciousness and temporal lobe integrity.

- It is difficult to express verbally but observable through behavioral patterns

- It is acquired gradually and improves with practice
- It does not usually require conscious effort (sometimes it does)
- Implicit and explicit are not completely separated. If the task is really hard, it’s better to leave
a bit of work to the explicit memory.
- It is little flexible and little modifiable
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 - It’s much more difficult to re-learn something than learning it from 0.
- It does not deteriorate much with aging
- It is phylogenetically older than explicit memories

Imitation is one way of implicit learning (if we mirror the behavior of someone else, since we’re activating
the circuitry), the physiological reactions after hearing an alarm, habituation as well, etc.

2.1. Non associative learning

- Produced by simple exposure to a stimulus. Just mere exposition already changes our way of
processing it and behaving.
- It is highly adaptive: it helps organizing behavior effectively

2.1.1. Habituation

Decrease in a reflex response to an innocuous stimulus presented repeatedly during a brief period of time.
It’s to a specific stimulus. Specific response.

2.1.2. Sensitization

Increase in a reflex response to moderate stimuli of different nature preceded by intense aversive stimuli.
Being exposed to the stimulus, if its context is aversive (the aversive context is the key) it can trigger a much
stronger reaction.

If you are frightened, a stimulus that in other situations might not startle you, will startle you. It’s not that
you associate it. It 's more general.

2.2. Priming

Can be perceptual, but it could also be cognitive. It facilitates the processing of specific materials to which
the subject has been previously familiarized to (that you have encountered already).

- Enables identification and cataloging objects and situations.

- If you had to buy something that you like and that you have been looking for models and
finally decide on one. Then, you start noticing that thing everywhere → you see it much faster
because you’ve familiarized with it (=priming).
- Enables memory formation after one exposure to the stimulus and can generalize to similar stimuli.

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 - Automatic, long-lasting, accelerates and increases the precision of the perception and
understanding of stimuli
- More effective within sensory modality but it also can generalize across modalities.

2.2.1. Perceptual

2.2.2. Semantic

2.3. Associative learning

Association between a stimulus and a response or between several stimuli

2.3.1. Classic conditioning

◆ Stimulus-to-stimulus. A neutral stimulus acquires the properties of a

relevant stimulus when they become associated.

◆ Ex: Pavlov

● A neuron disconnected to another one, when fire can’t make this neuron fire bc the
connection is not very strong. If the cell C is making B fire, then the connection A-B
gets stronger. At some point, while A can’t excite B, now, it will.

● At the beginning, there’s no association A-B. A (bell) is a cell in the associative cortex
which responds to a bell. But if Bell is bare with presenting food (C), the association
is form between A (bell) and B (salivary gland). The bell excites the salivary gland.

● Bell+food, now the bell is predicted and the digestive processes start.

◆ When associative learning is simultaneous we don’t need the hippocampus. But, if there’s a
delay between one and the other stimulus we need the hippocampus.

◆ Changes in the cerebellum and amygdala (if

aversive conditioning) → connexions

◆ Classic conditioning has been tried to use in

clinical chronic alcoholism. “Antabuse” is a
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 drug that metabolizes alcohol, and you become intoxicated very fast. With only a beer it does
feel very bad, and you associate this feeling with alcohol.

2.3.2. Instrumental (operant) conditioning

Stimulus-to-response. Implies learned responses. The consequence of a response determines the

probability in which that response will be reproduced (reinforcement or avoided (punishment). Enables
behaviour modification (Skinner).

➔ At the beginning you don’t know anything and explore the world, and this has consequences. You
need to learn these actions with that consequence.
➔ The reinforcement system will take place to make the association between the detection of this
stimulus and some behaviour, so we can receive some rewarding.

2.4. Procedural learning

Ability To Learn To Execute A Particular Skill (usually, but not always, involving motor actions)

➔ Skills can be motors, cognitive (how to position your

body) and sensory motor (sounds when you do
something)
➔ All of this is slow to learn, but is stable. There are so
pics.
➔ Changes in motor systems
➔ Skills are learnt through repeated execution→ we need to do it a lot of times
➔ Feedback is of the essence→ we need this feedback to feel that it’s good or
not.
➔ If accompanied by instruction and/or observation, accuracy increases→ if
there is a instruciontion is better because you don’t need to discover it by
yourself.
➔ Gradual learning: Fast at the start, slow when improving.

2.4.1. Motor skills

● Involve primary motor cortex, premotor regions, striate nucleus of the basal ganglia (caudate +
putamen) and cerebellum.
● During initial stages, usually strong involvement of prefrontal cortex and even
explicitation/verbalization (cognitive resources needed). E.g. Driving.
20
 ● The cerebellum compares what is intended with what happened
(feedback) and corrects to adjust movement.
● After practice, consolidation of specific cortico-striatal and cortical-
cerebellar loops enable AUTOMATION
○ At that moment we call Automatic all the behaviours
■ you don’t need any more rules and you become
faster. We aren’t conscious of doing that.

2.4.2. Cognitive skills

Prefrontal, sensory and motor cortices and basal ganglia are very important.

Ex: Parkinson disease affects substantia nigra, and is very important in processal learning. These people
can’t learn motor skills and they can’t do it.

Also dementia affects cognitive skills.

3. Explicit / declarative learning and memory

3.1. Introduction

➔ This memories requiere hippocampus and medial temporal lobe structures.

➔ Refer to general and Personal knowledge, declarative (able to be put in words), consciousness and
hippocampus (medial temporal lobe) dependent.
➔ Related with
◆ Semantic
● Learning: Ability to acquire information about facts of the world,
including ourselves, that we can share with our community.
● Memory: General information not tied to the context in which it was acquired.
Knowledge of the world, after exposure to episodic memory, the knowledge becomes
independent, when you don’t need the context anymore.

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 ◆ Episodic
● Learning: Ability to acquire information in a spatiotemporal context.
● Memory: Personal information context-related (space, time,relations...).
Autobiographical memories. When we remember an episode of our life.
◆ Spatial
● Learning: Ability to acquire information about one's environment
and spatial orientation.
● Memory: Information about navigation and cognitive maps.

3.2. Medial Temporal Lobe (hippocampus)

➔ Requiere hippocampus
◆ It looks like a seahorse. It is a continuation of
cortex that starts changing the citoarquitectonic
tissue, the type of cells in the tissue changes.
◆ Cornus ammonis → it’s called Hippocampus
trooper (?)
● CA1→ Cornus ammonis 1
● CA2→ Cornus ammonis 2
● …

3.3. Association cortex

Where is memory stored?

- In associative cortical areas (information storage)

➢ Prefrontal
○ Memory for motor actions
○ Reasoning
○ Language
○ Production
○ Working memory
○ Temporal sequencing of memory
○ Ex: names of people, faces, etc.
➢ Parieto-Occipito-Temporal
○ Memories acquired by senses
○ Semantic knowledge
○ Memories for spatial distribution of objects and people
○ Ex: visuo-spatial interaction
○ more related to perception than an action
➢ Temporal
○ Autobiographic memories
○ Spatial (navigation) memory

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3.4. Semantic memory

Semantic learning and association cortical areas

➔ PET study
◆ Task: Evoke people (social information), animal (information with organic things) and tool
names (information with inorganic things)
● information of
◆ Sample: Controls and people with cerebral lesions
◆ Results: Controls show different brain areas active according to semantic field. Patients show
specific impairments coinciding with lesions in those areas. There are differents activations
depending the semantic feel:
● People’s names→ Left Temporal Pole. This is heavily connected with inferior
prefrontal cortex throwing a fascicle. The information about association a person with
reproduction of your language throws for this.
○ People tongue problems (com el fenomen de la punta de la llengua, you
know?) → you need to say the name of someone or a thing and don't come
to you.
● Animals→ Left Inferior Temporal
● Tools→ Posterior Temporal and Parietal
○ Because we integrate information vision towards being able to execute the
action of interaction to an object.
➔ fMRI study (Hauk et al., 2004)
◆ Task: Passive reading of action words related to different body parts(e.g., kick, pick, lick).
● They show that when we read fastly a body part, they saw that these action words
would activate regions related to the region that would be used to do that movement.
● Reading would activate portions of the left motor cortex.
● We represent the concept of things related to how we process these things.
◆ Sample: Healthy adults
◆ Results: The referential meaning of action words has a correlation in the somatotopic
activation of motor and premotor cortex.

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3.5. Hippocampus and association cortex relation

● Association from many sensory areas

● There’s a loup (el cuadro en azul) → it’s the hippocampus formation → it’s necessary to consolidate
all memories of associations mostly.
○ All this information will go to the mamillary bodies of the hippocampus and this will go to the
thalamus (anterior nucleus) and then to the association cortex.

● The perirhinal cortex is the one who gets information about the object.
● Parahippocampal gyrus is more specialized for spatial information.

3.6. Two cortical systems for memory-guided behavior

There are 2 differences declarative memories and how and where they are encolled:

★ Anterior-Temporal System
○ Semantic memory
○ Recognizing familiar items
○ Multimodal object perception
★ Posterior-Medial System
○ Episodic (contextual) memory
○ Autobiographical memory
○ Spatial memory
○ Episodic simulation (imagining episodes)
○ Scene perception and spatial navigation
○ It’s encolled to all of this because it’s related to
the parietal cortex, regions for action, measuring
spaces and distances, etc.

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4. Implicit vs. explicit memory
● Implicit and Explicit memory usually interact, but their neural
substrate is different.
○ Implicit: Hippocampus independent.
○ Explicit:Hippocampus dependent.
● Weather forecast game → This is an experiment
○ The participants have to do a task that it’s called Weather
forecast.
■ Task: They have to predict how the weather would be
seeing a set of carts. When you predict if it’s going to
rain or if it’s sunny with a combination of carts.
■ The neuropsychologist takes a set of carts (each one
has different symbols) and asks you how the weather
would be (rain or sunny). The patient says: ou wtf how
I would know it? So, the neuropsychologist says: say something. The patient tries it
and say: it’s going to rain. Neuropsychologist: you’re wrong or, you’re 70% correct,
etc.
■ People improve in these tasks, but they don’t know why.
○ Implicit learning task (improvement with practice)
○ Parkinson patients: Do not learn, but remember what they did
■ They do not improve. When you ask 2h later they don’t remember.
○ Amnesiac patients: Learn but do not remember what they did
■ They do improve as well in these tasks.

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 5. Quantitative classification of memory
5.1. Introduction

➔ Immediate & short-term memory → These two make sense out of the world in the present moment.
If you are talking with someone and suddenly change the tone of the voice, it would be quite weird.
To be able to realise that he has changed their tone of the voice, you need a perceptual memory.
◆ Short duration (milliseconds to 15-20 minutes) → when we learn things
◆ Consolidation transfers it to long-term memory
➔ Long-term memory
◆ Long duration (permanent). It can be change, you can forget
➔ Working memory
◆ Ex: phone number
◆ Relates to attentional ability.
◆ Maintenance and temporal manipulation of information
◆ Needed in complex activities such as:
● Comprehension
● Reasoning
● Learning
➔ The overall reason
◆ Learning
● The painting is called The persistence of memory
◆ Immediate memory
● Immediately after, How is it called?
◆ Short-term memory
● 15 minutes after, How is it called?
◆ Long-term memory
● When finishing the lesson and within following days

If memory is not consolidated, we may forget in any of this types. We lose specific items on a category for
instance, but we can have a general category.

26
5.2. Working memory

➔ Neural substrates
◆ Prefrontal cortex in liaison with posterior
sensory and association areas (reverberating
activity)
◆ Working memory works through the
connections of the prefrontal cortex with
parietal regions mostly, but also temporal.

6. Synaptic plasticity
There is plasticity in neurons and also, there is plasticy in other structures like glia, that support neurons, but
are not neurons in fact.

6.1. Introduction

Modification of the neural substrate:

● Result of a change in conditions (experience). Changes in anatomical processes, ex: they learn
through modifying virtual connections. Experience will modify this neurosubstrates.
● During regular brain development or after damages/malformations (also in the brain!) These changes
in conditions can happen during the whole lifetime, since you’re developing until you die.
● Adaptive for survival and optimal functioning. These changes have functions to adapt you and make
you survive.

27
In brain organization, 3 types of processes interact, controlled by:

I. Genetic heritage (gene-driven processes): those processes that are related to genetic heritage, which
are related to phylogenetic changes in the neuro system, the experience that our species have in
evolution.
II. Common experience, expected for all members of a species (experience-expectant processes).
These are things that a particular individual of a specie is supposed to face. Every species, every
individual is supposed to face certain experiences, always. Ex: you’re supposed to get exposed to
the light, the sounds, the sensation of sucking on the lips, to have social contact… That’s extremely
crucial because whatever has been developed by genes phylogenetically, steps landscape to get
this common experience, and this common experience is going to shape appropriately for that
species and the neurosystem. → And from there the novel experiences that will be diferent for every
individual is what we call experience-dependent processes.
III. Novel experiences, different for each particular individual (experience-dependent processes).
Everything that is different in species and individuals. What that particular individual will learn in their
lives, will be different from every individual. We will relate to learning and memory.

EXPERIENCE-EXPECTANT PROCESSES:

● Regular experience expected for all individuals. Every individual in a particular species needs to
undergo/develop. The main idea: are those processes needed for an infant to develop properly, to
ascendal properly the neurocircuit that characterizes that species. Where adults would be facing
learning processes not development ones.
● Critical or “sensitive” periods. There are some periods in the development in which if experience
is not experienced, the particular event of development doesn’t take place. If the connections
happen during the critical period everything is well.

Ex: that’s the way cochlear implants work much better. If we implant them in the first years of life, that if we
do it in adult ages. In this way, the pobrish input which arrives at the cochlea is still able to connect the
sound with auditory structures (because kids are still in the developing phase).

● “Pruning” of overabundant synapses. When an infant is born, the brain birthing contains much
number of synaptic than an adult. WHY? It's difficult content, but basically, to keep only the important
connections, the ones that are necessary for survival. HOW? Kind of by random processes, some
connections become stronger and others weaken. But we don’t know the internal processes and how
the circuit works → he explains that but says that it is not relevant.
● BRAIN STRUCTURE: it would be impossible to have a heavily brain all interconnected, and also
wouldn’t be efficient at all. Apparently, the brain is connected as a social network, analogy with a
network of friends.
● Hebb’s rule: neurons that are fire together, wire together. Hebb's principle can be described as a
method of determining how to alter the weights between model neurons. The weight between two
neurons increases if the two neurons activate simultaneously, and reduces if they activate separately.

EXPERIENCE-DEPENDENT PROCESSES

● Adaptation to information unique to the individual.

● Do not take place during a stringent temporal interval. This does not take place in a particular period,
it can be extended in your all life.
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 ● Formation of new neuronal connections.

6.2. LTP & LTD

How do we strengthen or weaken these connections? Eric Kandel made a propose:

Strengthening and weakening changes are the ones that make we learn or we consolidate the learning in
memory. Learning and memory involve two main mechanisms:

● Long-term potentiation (LTP): stable and long-lasting increase of synaptic contacts (strengthening).
● Long-term depression (LTD): stable and long-lasting decrease of synaptic contacts (weakening).

The synapse (reminder):

● Communication between neurons uses synapses.

● The pre-synaptic neuron releases neurotransmitters
(NT) to the synaptic cleft.
• Excitatory: e.g. Glutamate • Inhibitory: e.g. GABA
● The post-synaptic neuron contains NT receptors at
the dendritic spines
● According to the NT and receptor types, a PEP
(post- synaptic excitatory potential) or a PIP (post-
synaptic inhibitory potential) will be
evoked at the post-synaptic neuron.

LTP: high pre-synaptic firing rate (reaches a membrane potential depolarization threshold at the post-
synaptic neuron determining the establishment of LTP). Discovered at the hippocampus by Bliss & Lomo
(1974).

It happens in several brain regions, such as: prefrontal cortex, piriform cortex, motor and visual cortices,
thalamus, amygdala…
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 ● Pre-synaptic neuron: high firing rate of escitatoru NTs (Glu).
● Post-synaptic neuron: depolarization by PEPs summation.
● Increase of the surface of the dendritic spine and increase in the number of receptors. → made
stronger and more efficient synapses.
1. The pre-synaptic neuron releases Glutamate.
2. The post-synaptic neuron contains receptors for glutamate.
a. NMDA: when glutamate is released, NMDA is blocked by Magnesium. So, that this blockage
is dependent on this type of depolarization of the cell. → That means: NMDA to be able to
open their channels need that the membrane potential it’s highly depolarizated, so it can let
the sodium in. The threshold for establishment of long-term potentiation, is the same voltage
that we need to separate magnesium from NMDA.

NMPA it’s a neurotransmitter, but also voltage dependent channel.

b. AMPA: are classic ion channels, that when glutamate is released, let the sodium go inside
through the cell. → And it depolarizes the cell.

AMPA are chemical, neurotransmitter.

c. Kainate
3. When 2 receptors are opened sodium can enter more fluidity, so depolarization can go stronger.
4. NMDA also leads calcium to enter the cell. When Calcium enters it will tell some storehouse that has
AMPA receptors: “let’s take some receptors from here and put them in the membrane” → so, make
the synaptic more efficient.

In this way, the amount of glutamate is able to excite more much the cell (because there are more
receptors now).

5. Calcium is also able to alter the genetic expression, so that more AMPA receptors can be made

The more we use a synapse, the stronger it gets. → the more glutamate, the more times that will bind the
receptors, and make more depolarizations.

6. Apparently, all this process triggers a message to the pre-synaptic neuron through nitrux-oxide (not
relevant). It tells to release more glutamate by positive feedback. So, that’s the way that synapses
get strengthened.

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LDP: low pre-synaptic firing rates.

6.3. Structural changes: synapses

We’ll see experiments in which we can see that numbers of synapsis change according to experiences.

Enrich conditions have signific high levels at certain days.

They see that correlates with having more synapses per

neuron than in impoverished conditions.

OLIGODENDROCYTES: part of SNC also changes with experiences. Oligodendrocytes are highly
specialized neural cells whose function is to myelinate central nervous system axons. So, in this
way, if there is more myelination, synapses are faster.

ASTROCYTES: have many functions, ex: modulate synapsis, providing

nutrients to the neuron, because the neuron is too busy doing synapsis.
→ So astrocytes feed the neuron.

The volume of astrocytes per cell increases when you increase the use
of the neuron. That means, the number of astrocytes will increase in the
brain where we are learning.

VASCULATURE: experiment: change in

vascularization, in macacs, you can see that those
that are running contain a higher level of
capillaries. Only during learning you will need this
vascularization, no persistence of the change in
time → same thing happens with astrocytes.

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 7. Emotional modulation of memory

7.1. Role of the Amygdala

Remembering stressing or emotionally intense situations is a highly adaptive ability

The amygdala:

● Has a key role in modulating memory consolidation

○ Potentiation
○ Inhibition
● It is activated by stress-related hormones (Adrenaline, Noradrenaline, glucocorticoids...) and evokes
an increase of consolidation through connections with the hippocampus, the basal ganglia, etc.,
modifying these regions plastically.
○ These regions contain receptors for and thus can be directly modified by these hormones as
well.
○ It is the stressing (activating) characteristics and not the emotional valence (nice or nasty) what
determines the participation of the amygdala.
○ Autobiographical and episodic memories are especially potentiated (albeit semantic memory
is also modulated).
● But, it can also cause inhibitory effects in several brain regions, affecting memory recall and working
memory (e.g. stay blank in an exam)

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7.2. Effects of chronic stress

TEMA 7: SPATIAL BEHAVIOR

1. Introduction

We have some systems that can provide with the information enough to be able to move in space and
interact with objects.

1.1. Definition of spatial behavior

Spatial behavior: any behavior that will allow directing the body through space (completely or partial).
- complete → translation (displacing ourselves → going from one position to another)
- partial → moving a part of our body in space
Any movement that allows us to displace ourselves or interact with our surroundings) almost everything we
do) is a spatial behavior.
Even thought processes → when we think about what we’ve done or when we’ve been (these are also
considered spatial (and time) behavior → we’re directing our thoughts in space and in time).

1.2. Subspace types

Sub-space types: useful classification in neurological

- Corporal → mediated by our somatosensory receptors without resolution enough to know where in
our body an object has touched us.
- Reaching → I interact with objects at a distance without the need to displace myself. Coordination
between the visual information and the motor movement (mediated by the parietal cortex).

33
 - Distal → I displace myself and get closer to reach an object and interact with them. It also allows us
to change the environment/space. We need to navigate to do this (know where we are, where do
we want to go, how do I get there, how do I get back, how long is it gonna take…).
- Temporal space → Thought process comes here. We can trace a path between episodes
(hippocampus) in time, which happened in space. There’s a time lapse when we move from one
place to another. In the brain the navigations in space and in time have the same core process, in
the hippocampus.

1.3. Allocentric vs. Egocentric frames of reference

We have to think what is our frame of reference (Where am I in respect to what? What’s the location of the
objects around my position? What’s the center of frame?)

Egocentric:
- If I take myself as the frame of reference, the computer changes location.
- It moves along with us (If I turn around, it won’t be in front of me but behind me).
- Ex: I’m in front of the computer. If I turn around, the computer will be behind me. The
computer is over the table, that’s in one room of my house that has a window. Doesn’t matter
where I am, the computer will be exactly in the same place
- Moraleja: If I consider myself as the frame of reference, the computer is changing
locations because I move the frame of reference with me in space (egocentric frame
of reference).
- In the reaching space and navigation.
- Important to interact with the objects. I don’t care where exactly is the object but we’re is it
in respect of me (is the coffee on my left? I don’t care if it’s in the table next to the corner, I
just want to know what I have to do to reach it).
- It extends even to those parts that are not of our body (ex: sword)
- PARIETAL.

Allocentric:
- If we center it on the surroundings or in objects.
- Matters the relationship of the object with cues of the scene.
- These cues help me know where I am.
- Ex: If the position of the door changed places in our room we would feel a bit
disoriented because we’re using the landmarks to position ourselves.
- Static, it doesn't depend on us.
- HIPPOCAMPUS.

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 2. Visuospatial processing

2.1. Two-streams model

There are two streams of visual processing that arise from not primary occipital cortex (the visual):

2.1.1. Dorsal

- Where and how pathway → vision for action

- Lesions towards the parietal inferior to frontal → impacts in spatial behavior.
- They know what the objects are but can’t direct the movement towards the object.
Also have a hard time moving along the space (orientation).
2.1.2. Ventral

- What pathway → recognize objects

- Lesions in the temporal inferior cortex→ agnosia
- They see but don’t recognize

2.2. Three-pathway dorsal model

What we know so far is that

the parietal where pathway
(ventral) is not unitary (not only
one occito-parietal pathway),
there are at least 3 different
pathways.

Hierarchically, the receptive

fields are much larger and the
information gets more
complex. Feeds from the
information of the visual and
some transformation.

35
- Occipito-parietal network
- Origin pathway
- From the occipital (visual) to the posterior parietal cortex
- Main job: Going from a retinotopic representation that changes to an egocentric
representation that doesn’t→ optic flow
- Even if we move the eyes/head, the retinotopic image changes, but we’re aware that
we are the ones moving and that our surrounding isn’t.
- Frame of reference: our body.
- People with lesions have a lot of trouble when they walk, and see the world moving as if they
weren’t the ones causing it.
- This divides in the 3 different pathways mentioned earlier →
- Parieto-prefrontal pathway
- Reaches the frontal eye fields (FEF) → controls eye movement
- If you stimulate the FEF electrically, we can see saccacs (the movement of the eyes -
also without movement as we’ll see).
- Premotor neurons control the voluntary movements of the eyes
- To direct the eyes and spatial attention
- There are different neurons that activate for specific directions of a movement of the eyes.
- We might be keeping our eyes still (no movement), but maybe our attention goes to
another particular position.
- Remembering where something was
- Allows to explore better spatially our world
- Parieto-premotor pathway
- Coordination of body and space representation
- Important for reaching movements
- Neurons that are not only sensitive to an object but to a relationship in space between
the object and me.
- Ex: Can I reach the glass? Yes. Then, a neuron will be firing to the glass since
it’s in my visual space. Since it’s not a retinotopic representation (it’s
egocentric), it doesn’t need to be where I’m looking at. If it’s not in my
reaching space the neuron won’t activate.
- Vision for action, interaction with the world in a proximal space
- Parieto-medial temporal pathway
- Goes towards the medium temporal lobe, and goes through the posterior cingulate cortex,
the retrosplenial cortex and the medial temporal lobe.
- Distal space and navigation as well as spatial memory.
- Combines the egocentric frames with the allocentric representations of space to be able to
navigate.
- Allocentric depends completely on the Hippocampus
- Egocentric depends on the Parietal Cortex

36
We know there are different pathways because lesions in different zones result in different outcomes or
limitations.

3. Positioning & Navigation

3.1. Allocentric vs. Egocentric navigation

Mostly distal space/objects and Hippocampus (and surroundings).

When we navigate, we usually combine both frames of reference (allocentric and egocentric) because when
we know the place very well, just by knowing one landmark we know how to get from one place to the other.
- Ex: in our house, if we want to get from the kitchen to our room, we know where to go, because we
have a map, which depends on the visual cues.
- Ex: if we’re in our city and we want to get to a certain store, we know how to get there when we exit
the subway, we might look at a map and check if we have to go left or right for how many streets,
we might recognize a building and we start moving.

We start moving because we have this cognitive map. We need these to be able to orient ourselves.

- Allocentric: It doesn’t matter exactly where I am or how do I move around myself, what matters is
these visual cues.
- Egocentric: If we were in a really foggy mountain in which we can barely see, we can’t rely on visual
cues to locate ourselves, we rely on information that we produce on our own while navigating
(egocentric information).
- Ex: I travel a distance that I estimate more or less depending on my speed, the time that’s
roughly gone by, then, I choose a direction even if there are no landmarks. If I want to go
back, I’ll undo this information.

Usually, when we are somewhere we don’t know, both function (and when we navigate in general, but mostly
somewhere unknown).

37
The moment you switch from egocentric information to allocentric, we start forming a cognitive map. We
navigate egocentrically in an unknown space and we start discovering landmarks.

3.2. 2004 Nobel prize in physiology

How do we know all of this?

- John O’Keefe
- May-Britt Moser and Edvard I.Moser

They discovered this positioning system of the brain, so they won the Nobel Prize of Psychology in 2014
(divided between the 3).

John O’Keefe (1939)

In 1971, while studying neuronal activity in the rodent hippocampus in freely moving animals searching for
food, O’Keefe described 8 neurons (out of 76) with a very particular response pattern.

- He talks about place cells (on the brain of a rat)

- Video 1: Recording with leds (the area lights up when the cell fires)
- Depending on the position of the rat in the box a neuron is firing
- Video 2: Also with multiple electros to check on the firing of multiple cells firing. The firing
draws a dot every time it happens.
- A rat in a maze
- The discharge of the neurons is different depending on where the animal is located
on the maze (different colors and codes)
- It doesn’t matter if the rat faces one direction or another (the egocentrism of the frame of
reference doesn’t matter; what matters is the allocentrism, that the animal knows where it is
in the scene).
- Video 3: They represent the abstract concept of the place (not the placing of something
discrete). Space representation.
- They have a receptive field of the place (n the hippocampus)
- Only fire when the animal is in a particular position in that space
- How do we know where we are? → these neurons represent the place where we are.
Basis of the cognitive maps.

38
With the tracking of the neurons and the combination of their firing we can know its position as well as
predict where the animal will go and where its been.

3.3. Place cells

Neuroscience provided represent spaces. A place cell

is a neuron which is mainly located in the
hippocampus but also the subiculum. They put
electrodes in the hippocampus rats in the subiculum
to register the activity to these neurons. The black
lines are the position of the animal, when it moves,
and in green is the firing of a neuron and it takes place
when the animal is in this portion of this place of the
box. The receptive field of a place cell is a portion of
a space, it depends on the animal's particular position
of the animal. They release that this could be the
bases of the cognitive map.

➔ Place cell receptive fields are selective to the location of the animal in a given environment (i.e.,
scene).
➔ Place cell receptive fields depend on the constellation of environmental sensory cues (mostly visual,
but also olfactory, tactile...) used as spatial cues.
◆ This selective location depends on sensory cues, use a spatial cue.
➔ If a spatial cue (i.e., landmark) is moved, or the lights are turned off, the firing pattern is
maintained→place cells respond to where the animal thinks it is located!
◆ If you are in the living room and you move your chair, you know that it’s your room and you
are there, and this map keeps it even if you have the eyes closed.
➔ Place cells form strong links to landmarks: if landmarks are changed unpredictably, place cells stop
using them; if landmarks define the space and they rotate, place cell receptive fields rotate along.
◆ If the window of your living room changes, you could think that it’s not your living room or
it’s a different place in the living room. That is because the place cells that orient you in the
scene establish strong links to landmarks and they define the space.
◆ A place cell will fire to a particular position of a space giving sense I know where I am,
depending on some important landmarks in space.
◆ But if you make landmarks that are unpredictable (change all the time), these cells stop using
them, because you will feel disoriented.
➔ The discharge pattern of a place cell is specific to a particular environment; a place cell can react
strongly to a location in an environment but exhibit no reaction at all in another one, or respond but
with a completely different receptive field (feature known as remapping).
◆ The particular receptive field of a place cell in an environment doesn’t apply to a different
environment. If 2 environments are very similar the place cells try to keep their receptive field
of position (the place cell would try to signal the same landmarks). But if the space is
completely new the place cell will signal that they are in that position and will be very
different.
➔ Place cell activity can be modulated by locomotion speed and/or direction.
39
 ◆ Depending on the speed and the direction place cell is modulated by other cues (some of
them of locomotion cues)

Exemples:

● Receptive Field Expansion When enlarging the scene.

○ Cover proportionally the same aspect of that, because it’s not a
new environment.
● Generation Of Secondary receptive field when introducing an object
partially dividing the scene.
● Receptive field reorienting as a function of a displaced visual landmark.
● Cell 1: Activity modulation as a function of the olfactory context (rate
remapping). Cell 2&3: receptive field modulation as a function of the
color or shape of the scene (global remapping).

Summarizing:

❖ A place cell is a neuron located predominantly at the hippocampus that represents a portion of
space, thus informing the animal about the position where it is located.
❖ This representation is flexible.
➢ The place cell rotates with that because that landmark is defining how you sense your position
in that environment. Place cells depend on the constellation of visual cues. When you change
the environment completely the mapping is
different.
■ When the room is black and white this
place cell rotes (like in the first photo of
place cells)
■ When the environment is completely
different the mapping is too. (like in the second photo of place cells)
❖ It implies the existence of a Cognitive Map.

3.4. Spatial memory & Hippocampus

In 1981, Richard G. Morris, in collaboration with John O’Keefe, developed his famous water maze.
Performing lesion studies they described a causal relation between the hippocampus and specific features
of spatial memory.

● If the water is clear the rat perfectly could see the platform
● If the water is clouded then the animal can’t use a visual cue, so it uses implicit memory. You leave
the animal at the same place of the pool
and it would find the platform, every trial
it would go faster and faster → This is
implicit memory.

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Radial maze

● Designed by David Olton et al. (1976) to study spatial working memory

● The Aim Is To Get Food As Efficiently As Possible (avoiding visit maze arms that do not contain food
or avoiding repeated visits)
○ Healthy rat:
■ The rat will explore all the arms once and eat all the food
○ Rat with a damage on the hippocampus:
■ It will visit all arms but only will repeatedly the arm in which it has already been. It will
not remember that it has been there, so there’s no food.
■ It will still implicitly learn to visit only these arms. It will keep on explore this arms
repeatedly

➢ Food storing birds (e.g. paridae, corvidae) hoard insects and seeds in the bark of trees, moss, etc. at
great distances, using distal landmarks not to repeat hiding places.
○
➢ These birds, unlike non-food storing birds, show a larger hippocampal volume.
○ Store food in different places and these birds have a seasonal behaviour. So, the
hippocampus grows physically in these seasons when birds store the food in different places,
depending how much food is stored in many different places.
➢ The size of the hippocampus varies seasonally (due to neurogenesis), probably as a consequence of
its involvement in behavior and cognition related to storing and retrieving food.
➢ What effects do you think a lesion in the hippocampus will have in
these birds?
● Taxi drivers needs to know the shortest route, and it their
hippocampus is larger than the bus drivers (always do the same route)
○ Right posterior hippocampal gray matter volume is larger in
taxi drivers than in controls.
● Topographic memory
○ The right posterior hippocampal is more active in this type of
tasks

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 - In this study they have 2 types of tasks:
- Related to spatial memory remembering where is some place to go.
- Related to episodic memory → ex: “what item where do you pick
up from the library”
- Several studies using fMRI and virtual reality (ecological
approximation) show increased activation of the hippocampus. In
particular:
- Right hippocampus to spatial memory → is more activated
- Left hippocampus to episodic memory

3.5. What do we need to navigate?

The discovery of the place cells and their firing patterns supported the idea that position is represented in
a cognitive map, probably located at the hippocampus as anatomical, functional and lesion studies in
humans and animals suggest.

However, in order to navigate, what else do we need to know?

➔ Position
➔ Direction
➔ Measurement of travelled distance
◆ also to know how much we travel.

We need to have all this information

3.6. Head-direction cells

In 1984, James Ranck discovered some neurons that fired when the rodent
pointed its head towards a particular direction,independently of its position.

➔ He found a particular type of neuron that fired depending on the head points. Describe where
neurons that fired anywhere in space but only in the head points to a particular direction!!!
➔ The head directions cells also rotated with a landmark very much at the place cells. These cells (some
of them) depended on the direction of the eyes but many of them depended on the direction of the
head. Sometimes completely independently of the direction of the eyes, sometimes modulated by
the direction of the eye, but dependent on the direction on the
head.
➔ Receptive field of a head direction cells and the name indicates
selective of a particular head direction independently of the
spatial location of the animal.
➔ Head-direction cells receptive fields are selective to a particular
head direction, independently of the spatial location of the
animal.
➔ Head-direction cells receptive fields depend on the constellation
of environmental sensory cues (mostly visual, but also olfactory, tactile...) used as spatial cues.

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 ➔ Each cell exhibits a preference for a particular direction, but all of them
rotate coherently along with rotation of spatial cues.
➔ They are located predominantly at the subiculum, but they exist in each
structure of the Papez circuit (hippocampal formation (subiculum) → fornix
→ mammillary
bodies → mammillothalamic tract → anterior thalamic nucleus →
cingulum → entorhinal cortex → hippocampal formation).
➔ Head-direction cells provide the animal with a sense of direction as well
as a direction reference for the rest of spatial representation forms at the
hippocampus.

3.7. 2004 Nobel prize in physiology (bis)

Positioning & Navigation, 2004 Nobel Prize in Physiology (bis)

In 2004, Moser & Moser recorded neurons from the medial entorhinal cortex (mEC), given the connectivity
of this region with the subiculum (head-direction cells) and the dorsal hippocampus (place cells and spatial
memory), and found a very, very peculiar response pattern...

We were talking about how the Moser & Moser (that basically met in a lab that was collaborating with Morris).
They were very good at doing mini-slabitions, so many precise lesions of the hippocampus, and they started
discovering that the posterior part of the hippocampus (the dorsal aspect), was more related to spatial
memory. And then, they started to learn with John O'Keefe, about doing the measurements with electrodes,
and they pointed to the ento-neocortex. They found some neurons that have a very particular type of
response. In which at the beginning it could look like they were sort of place cells.

Basically, one of the post-docs (post doctorados), discovered that these neurons had a particular way of
locating in which the receptive fields, the pick of the response, was separated by a regular distance. So, they
form like a matrix. → that’s why we call them GRID CELLS.

3.8. Grid cells

Grid cells

● Grid cells receptive fields are multi-peaked. This pattern of response in which the vertices of a
triangle.
● The organization of these peaks is highly regular and precise, with maxima at equilateral triangle,
covering the whole scene where the animal is placed even if it moves erratically in it. It doesn’t matter
how you move or what you do, it will keep signaling like a tactic distance.
● Firing patterns of anatomically close grid cells exhibit similar orientations and scales, but vary in their
degree of spatial offset. In other words, interestingly, there is an anatomical arrangement of the grid
cells of the property of responses: those that have the location close anatomically, what you can see
is that they find patterns that have similar orientations.
● There are several properties that you can have:
○ Grid spacing: the distance between the activating peaks is short or long. It means that if I
advance 1m, they will fire again, and then 0.5m doesn’t, but at 1m it fires again. Or maybe
only fires every 5 meters.

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 ■ The spacing in anterior-posterior or ventral-dorsal axes changes being the more
ventral portions of the entorhinal cortex with largest spacing, and those more dorsal
more shorter as step sizes.
■ Dorsal are close, have different offset, but similar orientation and similar spacing. And
from dorsal to ventral toy make the spacing larger

○ Shifting orientation: it could shift, so that this grid cell can be shifted in orientation.
○ And, they can be shifted in offset, shift laterally → they do not coincide, they do not rotate,
they do not change spacing, it’s just an offset .
● So, the thing is that anatomically closed grid cells have very similar scale, so spacing and orientation.
But they vary in offset.
● Together, all grid cells are organized in a topographical, crystalline structure in discrete stepwise
increments of grid spacing from dorsal to ventral.
● In no proper words, this is a frame in the brain to measure reality.
● The orientation of the grid firing pattern reflects the scene where the animal is located in, but the
variation is completely coherent (unlike in place cells, which was decolorated) and, if landmark
changes are minimal, it does not vary (unlike in place cells, grid cells do not show remapping and
always respond to all scenes).

Summary:

● A grid cell is a neuron, predominantly located at the medial entorhinal cortex (mEC), that shows a
spatial response pattern distributed in a precise and regular fashion (geometrically in equilateral
triangles).
● This representation is generally invariable to changes in the scene, albeit these changes affect its
orientation.
● Imply an abstract representation of location, like place cells, but also represents the application of
an internally generated cognitive frame, with a precise and regular structure, to the outside world,
allegedly to measure distances between landmarks. → the trick part.
● Therefore, they fall in between allocentric and egocentric representations of the spatial scene. It
helps measure the travelled distance.

4. Spatial representation vs. episodic memory

The combinatorial capacity of the space representation systems (place, head-direction, grid cells) is so huge,
that it would be surprising if it was only developed to navigate.

The main idea is that the combinatorial aspect of all the previous mechanisms is so huge, it’s a complex and
expensive system that we use to navigate in space. People started to think we us have a mechanism such
expensive, complex… if animals can move around and navigate in space without all these mechanisms.

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 Ex: typical example, is the migration of monarch butterflies (they migrate from Mexico to Canada). It
takes them a lot of time, and they do it in 4 generations → so navigation is indeed on their genetic codes.
Insects are able to navigate with much simpler neural networks, why would we need this very complicated
system.

It has been suggested that the same mechanisms used to define position and spatial relations in a map can
be used to symbolize events, objects and living beings. In other words, scientifics say that this mechanism
which represents and symbolizes the space and spatial relations, can actually be used to symbolize events,
objects, living beings, people, things that we have done, things that we will do…

And that instead of being related in space they are related in time, so here, there is when Georgy Busaky,
if the brain would care about memory as being something different than spatial navigation, if it would care
about representing time and space differently → then, this 2 things would have 2 different dedicated
systems, but they don’t. IT’S THE HIPPOCAMPUS. Ex: take a person with Alzheimer, they don’t know what
they have eaten, and they get lost. Same way as in spatial memory 1 route goes at: get out of my house, go
to the coffee shop… you can do the same with memory: when I woke up I did some exercise, then I had my
breakfast….

The thing is that I remember the sequence of an event, a day… in the same way as I remember the way in
a route.

Because the main thing is that this mechanism just cares abouts taking in information, giving it a
representation and linking it according to what comes first and what comes next. → In other words,
repeated exposure to the same scene generates the representation of the space independently of temporal
context. Likewise, repeated exposure to experiences events can generate semantic memory. Maybe for
egocentric navigation to allocentric navigation is the very same thing as in episodic memory (which is
reference to yourself that at some point gets separated to the sequences and just get the representation of
the things such as the landmarks in a map) to semantic memory (first there was a context, then there is no
context).

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 TEMA 8: LANGUAGE
1. Introduction

1.1. Linguistic family tree

There are supposedly around 7.000 languages in the world and there is no known human community (actual
or past) without speech. There are as many kinds of speech as human communities.

1.2. Communication vs. Language

Is language communication? Yes, language is a form of communication used for humans, but not all ways
of communication are languages. Then, what does “to communicate” mean?

➔ It is an action performed by an emitter that alters the behavior of a receptor. Implies mutual benefits
across individuals (we could discuss this part).
◆ It’s a really general definition.

Communication pathways:

- Chemical: Odors, pheromones, within our bodies (hormones, NTs, insuline…)

- The most prevalent
- Visual: Emotional facial expressions; dances; characteristic movements
- Auditory: Bird, whale and dolphin “songs”; human speech
- In humans, the major communication pathway is auditory (oral)
- Can be learnt (the songs of the birds, for example)
- Tactile: Grooming

What is language?

➔ Any entity has a form of communication so, what makes language

special?
➔ A code, a tool to communicate complex information, to transmit
ideas (emotions, intentions, desires) and to organize thought
(connect ideas, make plans, decisions, predict…).
➔ It allows associating arbitrary symbols (sounds) to specific
meanings.
◆ It requires symbolic abilities, which several animals have.
➔ Language exhibits qualities that go beyond this communication with arbitrary symbols
◆ It exhibits qualities of productivity and displacement.
● Productivity enables using rules (grammar) to make new symbols (not arbitrary, it
follows rules).
○ Ex: someday Youtube was put on the internet and people started uploading
videos. Then, people started working uploading videos -> Youtubers.
● Displacement is that you can use these symbols to talk about things that are not
present at the moment.
➔ All human languages (oral, written or gestural) have the same components.

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Do animals use languages?

- Non-human primates use sounds, gestures and facial expressions to communicate danger, territorial
claims, etc.
- Darwin (1871) realized the parallelism between the learning of songbirds and human speech.
- Apparently, brain regions analogous to humans are used (perceptual and motor)
- Bees, dolphins, chimps… do they have a rudimentary form of language?
- Dolphin communication is very complex, at least as complex as human communication (they
might also have productivity and displacement)
- We have to find the actual meaning, so it’s hard
- Bees also have certain dances to communicate several things regarding the placement and
quality of food, which is complex as well
- What about dogs?
- They are a human invention, but they are very different to us.
- There are asymmetries in lateralization.
- Words are mostly processed by the left hemisphere (fast signals), while most processes are
in the right hemisphere (slower modulation).
- They don’t use language the same way that we do, but there’s the asymmetry in the brain so
there’s probably an underlying more basic mechanism.
- Intonation patterns are important (right hemisphere) .

Therefore, albeit as humans we exhibit…

- An extensive vocabulary
- A great syntactic complexity
- A high combinatorial level
- Well developed neural pathways to process linguistic information

… It is difficult to distinguish the frontier between the concepts of communication and language and,
probably, animals like dolphins use a language system with a level of complexity similar to ours! We just
don’t know much about others.

1.3. Language components

- Phonemes: Minimal unit able to change meaning (ex. Road vs. toad)
- Morphemes: Combination of phonemes. Minimal units constituting words (ex. Ta-ble).
- Lexicon: Set of words in a language (50.000 in adults)
- Syntagma: Admissible combinations of words to form phrases.
- Semantics: Meaning of words and phrases.
- Prosody: Vocal intonation able to modify the literal meaning of words and phrases.
- Discourse: Set of phrases forming a narrative unit.
- Pragmatics: Communicative intention, based on the expected effect produced in the receiver.

1.4. Language processing levels

Processing levels involved in language comprehension and production.

We say that the tiger is approaching - We’re producing a sound - There’s a phonetic and acoustical analysis
- It will contact with a phonological (level of the sounds/characticals) presentation - Combination of those -

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 Connection with mental-lexicon storehouse
(we’re we store words) - Access to the meaning
and grammatical codes - Access to the
concepts.

(To speak à ) Concepts should be able to

connect to the mental-lexicon to be able then to
select the words or ways of saying things. Then,
they should be translated into different
phonological representations, which then need
articulation processes.

1.5. Human language characteristics

Speech is a primary ability of language because it’s innate, we’re born with the ability to acquire speech.

The secondary abilities are not biologically determined, although there are biological mechanisms that will
be used to learn them, but they are culturally determined.

- Writing and reading

- They are an idea that someone had at some point and then, since it was useful, it went
through generations.
- We need someone to teach us.

2. Human language development

2.1. Universal steps

These steps are common in every child, wherever it is.

As we grow older we lose the ability to speak any language

so that we become experts in our own, but, at the beginning,
we’re all able to speak in any language.

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2.2. Development in babies

- Babies prefer speech sounds vs. other sounds (specifically the voice of its mother)
- During the first months, babies discriminate speech in different languages and speech from inverted
speech (grammatical comprehension, they can pick up some statistical rules in languages).
- Inverted speech = speech backwards, which sounds unnatural
- Phonemic perception: From 0 to 2 years old, babies can discriminate phonemes from all languages
in the world, but when they acquire their mother tongue, they become selective for its phonemes.
- Ex. Japanese babies > 10 months do not differentiate [r] from [l] (Werker et al., 2005), but if
they were exposed to english they would recover the ability within that sensitive 2-year
period.
- After the 2 years, we’re starting to be out of the sensibility period.

Last trimester:

- End of migration.
- Boost in connectivity, gyrification of the brain, synaptogenesis, myelination, neuronal differentiation
and glia reproduction
- Determined -> when we start being exposed to language
- more grey matter in regions that we’ll use to perceive and to act in simple forms. For instance,
the first thing to mature in the grey matter are primary areas and motor areas. Later on, these
will mature more. Then, those more related with language (like the broca area). Then the C
shape (last picture - dorsal and ventral pathways of speech).
- Not a linear process.
- The growth of the brain goes according to the universal steps in language acquisition.
- Goes in line with the exponential expansion of the vocabulary.

At 6 weeks old, long-range associative fibers are already similar to adult ones.

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3. Lateralization

There’s a few evidences regarding the differences between hemispheres:

❖ Structural
➢ Sylvian Fissure
■ (from the temporal lobe to the frontal and to the parietal).
■ Longer in the left hemisphere
● This means that probably the regions contained are larger too.
■ The angle is different, more closed
➢ Planum Temporale
■ Inside we have the primary auditory regions, to be concrete in the Heschl's gyrus.
■ This is where men grow grey hairs before (signaling aging)
■ Happens on right handed people
● L>R -> 65% adults
● R>L -> 10% adults
● women > men
● Evidence of asymmetry from 31st week of gestation
❖ Functional
➢ Neurological patients
■ Surgical interventions: Callosotomy
● Patients had epilepsy and they didn’t want it to spread to both hemispheres.
● Callosotomy = to cut (the corpus callosum, since it’s the most important
highway that connects both hemispheres)
● Sperry -> tachistoscopic presentation
◆ tachi = fast
50
 ◆ toscopic =visual
◆ Presenting images really fast, in a way there’s barely any time to
recognize it. He observed that when you present a very fast image to
one hemifield of vision, if the hemifield was the left, and ask the subject
what they saw, they cannot report it. If you present it to the right
hemifield they can (since it’s processed by the left hemisphere).
➢ This means that the language is on the left (not exactly, because
when you present it to the left hemifield, even though the
subject cannot name it, they can still recognize what they saw).
➢ They access the information and can express it in different ways,
just not verbal.
➢ This led people to think that the production abilities of
language are lateralized to the left hemisphere.
■ Wada test/intracranial electrical stimulation
➢ Typical people
■ Functional neuroimage
● In babies there’s also more activation of the left than the right. If there’s this
left dominance for linguistic processes, what does the right hemisphere do?
■ Dichotic listening
■ Tachistoscopic presentation
■ Tapping

3.1. Functions of the right hemisphere

❖ Lexical & Semantic processing

➢ People with RH lesions perform worse in:
■ Lexical & semantic discrimination tasks
● (words and their meanings)
● It can alter the way we see when we
speak
● Point to the picture showing what
the listened or read word means.
■ Denomination tasks
● Say what an image is
❖ Pragmatics
➢ People with RH lesions exhibit difficulties in understanding metaphors, irony, sarcasm or
humor.
■ Making use of the context.
● Might be cultural as well.
❖ Prosody
➢ Rhythmic, emphatic and melodic aspects of speech. Related to musical ability and emotion
recognition.
■ Difference between affirmative sentences and questions (more relation with the right
hemisphere than the left).

Beware trivializing the left-right hemisphere dichotomy!

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 - There is no such thing as left=analytical and right=creative
- Anatomic asymmetries and functional localization of linguistic function to different hemispheres
exists, but it is not that simple and depends wholly on how we operationalize what we consider a
particular function different from another.

4. Speech comprehension & production

4.1 Approaches to study the neural bases of language

The first authors reporting changes in behavioural related

to language abilities and the relation with functions.
Defining function and putting labels is not trivial. We are
going to explain these approaches...

4.2 Localizationism

❖ Broca
○ Paul Broca (1824-1880)
■ He was a french neurologist. He describes several cases that have problems in
producing speech, in talking.
● 1863. Publishes a study of 8 cases with left frontal lesions and language alterations.
● Leborgne (Monsieur TAN), 51 years old, presenting afemia (inability to speak) from his 30s. Normal
comprehension.
○
○ Leborgne has an inability to speak because of these lesions (area de Broca), and even though
he had relatively normal comprehension, he couldn’t speak. When he did speak, mostly there
was only one word that came out of his mouth. This word was "TAN".
● Lelong, 84 years old, reduced language production
ability (only 5 words).
● These people had lesions at the frontal cortex in the
left hemisphere in a region that is the inferior frontal
gyrus.
● Post-mortem analysis of the patient “TAN” brain
shows a lesion at the left 3rd frontal
gyrus→Brodmann area 44 (aka. Broca’s area).
○ He would study the composition of the brain, the tissue, and for this reason, he divided the
brain in a lot of different areas.
❖ Wernicke
➢ Carl Werincke (1824-1880)
■ German neurologist
➢ He had patients with different types of symptomatologies, and they couldn’t understand what
they were being told, speech, but could produce speech.

52
 ■ You would ask them something and they would
answer you something completely different. They
could understand the meaning of the words. They
would talk fluidly, but without sense and with a lot
of errors.
➢ 1874. A patient with speech
comprehension problems but with fluid
production, albeit with errors.
➢ The autopsy revealed a lesion at the left superior temporal gyrus→ Brodmann area 22 (aka.
Wernicke's area).
■ Studies post mortem would reveal that the lesion were mostly located at the posterior
part of superior temporal gyrus.
➢ Wenicke concludes that this region is related to the auditory storage of words and that it
would be connected to Broca’s area (to be able to produce the intended words)
❖ Dejerine
➢ Jules Dejerine (1849 - 1917)
■ French neurologist
➢ Describes a key region for literacy (reading
and writing)
■ He describes functions of the people
that have lesions in the angular
gyrus. They have problems with
reading and writing.
➢ Brodman’s area 39 Left angular gyrus

4.3 Classic Connectionism

❖ Lichtheim
➢ Ludwig Lichteim (1845 - 1928)
■ German physician
➢ 1885. First psycholinguistic model: The house
■ Several parts of the brain should contain several functions
that would be interconnected to support language
function.
■ If the input is auditory and we perceive it with our ears then has to be sent to the
auditory representation of words. This Auditory representation should be connected
to a center of concepts.
● Ex: Car
◆ It has to be a representation of the word and has to connect to the
meaning of this word. It’s be able to comprehend speech. And then,
find the motor representation
◆ You should go to the conceptual center, meaning of the car and find
the appropriate representation in the motor domain. Your tongue,
mouth, etc. are coordinated to produce this word.

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 ■ B: Conceptual center
■ A: Auditory representation
■ M: Motor representation
■ Inputs
❖ Wernicke-Geschwind model
➢ 1967. Norman Geschwind extends Wernicke’s proposal.
➢ Norman Geschwind (1926 - 1984)
■ American neurologist
➢ The Wernicke-Lichteim-Geschwind model appears
➢ Key elements in the model:
■ Broca’s area
■ Wenicke’s area
■ Arcuate fasciculus (axon bundle connecting both regions)
● White matter tract
➢ Task example:
■ Word repetition
1. Auditory cortex: Sound coding.
○ To hear sounds
2. Wernicke’s area: Word identification.
○ Comprehension area, you identify the word.
3. Arcuate fasciculus: Information transfer.
4. Broca’s area: Speech motor code.
○ To move coordinately
5. Motorcortex: Motor control of the phonetic apparatus.
● Word reading (transformation of the written code to a phonetic code):
1. Visual cortex: Vision coding.
2. Angulargyrus (at theoccipito- parieto-temporal region):
Code transformation to a similar activation that produced by listened speech.

Model prediction: What would be the behavioral consequences of a lesion at the arcuate fasciculus?

- If you would lesion the arcuate, you will have what is known conductuonal X. You will preserve
comprehension, preserve promotions (so you will be fluid), but due to this disconnection there’s no
ability to repeat words, you can’t connect very well the storage of the word. You will be very fluid
but paraphasic.

Besides, there’s the need of a connection between those brain areas and other associative cortical areas
(frontal, occipito-parieto-temporal and inferior temporal) for a correct language production and
comprehension.

➔ Memoric information is distributed to all cortex, mostly in

associative areas, but is distributed anywhere. It has to be a
connection of these regions holding the concept of the word.

Connectivity Lesions→Expressive And Receptive Language Disorders

(paraphasia, lexical confusion within a semantic field...)

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4.4 Contemporary Connectionism

Contemporary Connectionism is Motivated by:

● Technical improvement: Precise neuroimaging, precise lesion studies.

○ Brought better definition of lesions, and functional in techniques, more precise ideas on
language function.
● Conceptual improvement: Better delimitation of language functions and its constituents.

Language is not a product of concrete brain regions

It is the result of synchronized activity within large neural networks, constituted by cortical and subcortical
regions and their connecting pathways

For instance, Broca’s area is much more complex than initially thought.

Using innovative anatomical techniques (cytoarchitecture analysis, immunocytochemistry) we now know that
Broca’s area counts with 10 or more sub- regions.

Presumably, each of these participates in different functions:

➢ Working memory
➢ Segmentation and union of phonetic and syntactic information
➢ Linking phonemic sequences with motor gestures
➢ So, Broca may not be the site of articulation, but a key node in manipulating and forwarding
information across large cortical networks responsible for key components speech production.

Major conceptual advancement. Dissociation between:

● Speech perception: Linguistic processing in sublexical tasks, such as syllable discrimination.

● Speech recognition: Set of computations transforming acoustic signals in neural representations
contacting the mental lexicon.

Greg Hickok - David Poeppel → They are the authors of this dissociation.

➔ You can’t study processes that are for instance sublexical (for example, discriminating syllables) with
studies of speech recognition, because speech recognition is level higher.

55
The first stage when auditory input enters in the auditory system, which is a computation of the acoustic
signal by the superior temporal gyrus, which is the initial analysis of the acoustic material, hearing. It has to
“analyse”. The main idea is that the tympanic membrane fluctuations transmitted as electrical signal to the
auditory trial have to represent the composed connected with other regions, so that the acoustic material
incell is processed. This is connected with the superior temporal sulcus.

From Mid-post STS (Phonological network) emerge 2 pathways (dorsal and ventral pathway).

From here, there will be 2 streams: one that goes up/ parietal to frontal, one that goes down/ towards
inferior temporal and at the end connects with frontal. So it can be defined as, one pathway dorsal, and one
ventral.

● Ventral pathway:

As in vision, the ventral pathway relates to meaning (it’s called the WHAT pathway). In the ventral pathway
of language, these regions participate in ascribing these phonological representations (that you have in the
phonological network), in meaning.

See that it is mostly bilateral. → This means that you need lesions in both hemispheres in order to lose your
ability to recognize concepts from oral language.

It entails the Medial Temporal Gyrus (MTG) and the Inferior Temporal Sulcus (ITS). This is involved in speech
recognition.

● Dorsal pathway:

Language for action, so production of speech. When you go from this fronto-temporal analysis and
phonological network towards the dorsal, you go through a very interesting region on the most posterior
part of the sylvian tissue, that’s called area SPT (sylvian-parieto-temporal).

In this region, SPT, you can find neurons that have audio and motor properties. So they would be active for
instance to the characteristics of a sound, like rrr, but also when you produce this sound and do this
56
movement it could also get active. So, the neurons hold up the representation that it’s mixed between how
something sounds and how we make the motor part. Here, these neurons are getting auditory info (from
very complex speech sounds) and motor info.

SPT has reciprocal connections with the phonological networks and the spectro-temporal one.

You are producing before you’re activating this network. At the same time you are listening. So, the activity
gets messy at some point. If comprehension and production were completely separate we would not have
these missmatching moments/confusions.

See that the articulatory network is lateralized. So when we say that language is lateralized as a function, be
careful, not everything is lateralized. Production in most of the people is lateralized. But not all language
functions.

Say something and write something different. It's so hard to do, it takes a lot of effort.

● Hickok & Poeppel Dual Route model (2004) is anatomically supported as well by white matter
tractography.

These 2 networks are supported by the connectivity that links all these regions together. But we can see
how the dorsal pathway and ventral pathway are supported by different fasciculus.

○ Arcuate fasciculus, it links these regions from the temporal low towards the anterior.
○ With fibers in the ventral circuits, another fasciculus → it connects the posterior and anterior
regions of the temporal with also the frontal part.

5. Neural bases of literacy

Reading

When you don’t know a word, you have to go letter by letter decoding into sound, this is slow, it needs
time, but it’s quite precise. The longer the word the more time you spend.

Instead, in words that you already know, you have a representation of the global image of the word. So
when you see technology, that is a word with many letters in comparison to sun, but you read both of them
equally fast. Why? Because you have the global picture of the word, and that goes through a different route.
This one is fast, but not precise, because in the example it was TECHNLOGY, but you read TECHNOLOGY.
You don’t see that the O was missing. That’s why we make orthographic mistakes although we revise the
essay.

So we have 2 pathways:

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The global pathway it’s called lexical pathway, when you get the whole word: you see the word and
recognize the drawing.

Instead, when you have to decoding graphemes (the little drawings of letters into sounds) into phonemes,
we are using the phonological pathway. We use it for unknown words, things that look like words but they
are not (like pseudowords). This is the pathway which is really affected in dyslexia.

During the process of learning to read, kids start with the phonological pathway (more precise but slower).
Progressively the use of the lexical pathway increases and kids become faster readers.

We know that the neural network supports this reading ability, these systems of reading are different.

When we go with the lexical pathway, global word recognition, we are activating mostly the visual ventral
pathway. So, we are activating a lot the fusiform gyrus which is specialized in processing complex visual
information, like faces, and regarding the information we see as a whole. This would be the ventral pathway
for reading which tends to dominate more

And, instead when you go through the phonological pathway, letter recognition, where you have to be
using visual cortex to see the letters and then go letter by letter decoding them into a sound. In this, seems
more important the angular gyrus. This would be the dorsal pathway for reading.

Experiment:

An fMRI study with Japanese readers supported the role of the ventral and dorsal pathways in global and
phonological reading, respectively.

Japanese uses two different symbolic codes:

● Kanji: symbols representing concepts.

○ Ventral visual pathway (global reading).
○ Visual cortex → bilateral (left-dominant) fusiform area (inferior temporal gyrus)
● Kana: phonetic representation of syllables, codes acoustic information.
○ Dorsal visual pathway and Broca’s area.

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 ○ Visual cortex → left supramarginal gyrus (SMG [Parieto Occipito Temporal Area], close to the
angular gyrus) & left inferior frontal gyrus (IFG: Broca’s area).
● Both pathways will end up connected to the associative cortical regions related to word meaning.

Writing

Writing involves several cognitive processes:

1. Read (or think) words: for that specific meaning we want to express, we must find the specific word,
decode the phonemes that form it and convert them into graphemes. Orthographic correction needs
this process as well. You can read globally, but you cannot write globally.
2. Translate graphemes to movement: be it on a keyboard or by hand, we need to plan to plan a
specific sequence of movements.
3. Move fingers to actually write. Have motor areas related to the properly action.

Neural bases of writing involve:

● Linguistic regions: perisylvian areas (inferior and posterior temporal), ventral premotor cortex.
● Motor regions: primary motor cortex, somatosensory cortex, premotor cortex, thalamus, putamen
(basal ganglia).
● Visual regions: dorsal parietal lobule.
● Regions specific to writing: medial and superior frontal gyrus, superior parietal and cerebellum.

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 TEMA 9: NUMERICAL PROCESSING AND CALCULS
1. Introduction

We are surrounded by numbers. Numerical abilities are key in our daily lives. → We need numerical abilities
to understand day-by-day activities, although we may not realize how important they are.

The way we think of them: we usually think of the graphical representation, but there are so much deeper
meanings.

Dyscalculia, sort of dyslexia but only in numbers. It can go deeper on problems than dyslexia.

2. Concept of magnitude
2.1. Magnitude vs. quantity
● Magnitude: the property that allows us to measure. In other words, a property that all objects,
phenomena and relations between them possess, allowing their measurement. That measure,
represented as quantity, can be expressed using numbers on the basis of comparing it to another
object or phenomena that is taken as a pattern.
● Quantity: quantification of magnitude in any system of units. In other words, quantity is the measure
of a magnitude.
○ Any quantity is defined by comparison: somebody decided to do it in this way, but it could
be different.
■ Ex: we can measure temperature (magnitude) in different types of quantities, like:
degrees, fahrenheit… but temperature still being the same, although we quantified it
differently.

For instance: time is a magnitude, but 12h is a quantity.

2.2. Non-symbolic vs. representation

The main point is that magnitude can take 3 different forms in our processing of them:

● Non-symbolic
○ Uses a continuous representation of elements. Not discrete.
○ Allows distinguish between a lot and a little; more or less; little and big… You don’t have to
count, ex: to know which tree has more apples.
○ This ability is based on Weber's law:
■ to be able to perceptually detect the change, it doesn't
matter the absolute value, it matters the differential value.
In other words, the change in intensity of a stimulus
necessary for the organism to detect that change is
proportionally constant to the intensity of the original
stimulus, instead of a constant quantity.

● Symbolic
○ Uses a discrete representation, arbitrary assigned to a code.
○ Basically, allows counting.

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Experiment: domination of the quantity of elements (Mundurukú)

This experiment tends to prove that although symbolic representation it’s important, but not absolutely
necessary. Their language does not have numbers to represent numbers from 1 to 5.

The first thing they did, show them on a screen different sets of dots. And they ask them how many dots
there are, and they response what it is reflected on the graph:

Una vez que superaron los 5 dots (de 5 a 10), en general decían one
hand, some/ not many. Y si había más pues usaban many or really
many, pero no contaban de manera discreta después del 5.
So, scientifics saw that discrete arithmetic abilities, like 5+4, would
require language in a way that even when trying to make these
additions with numbers that they do have words, like 2+2+3+4+1,
they were very bad at doing that (Mundurukú). So, everything that was
described arithmetically, they were not good at all. Compared to
french people that could do it much better.

But when you would ask them to compare sets of numbers, ex: which one has more dots. Here you will be
using the non-symbolic ability. So here, we can see how french and mundurukú population, they will perform
the same, following Weber's law. There is almost no difference in proportion.

Performance in symbolic numerical tasks

Symbolic representation depends on
language and education. As you can see how
it increases with the grade of education.

3. Development of numerical abilities

Language is only necessary to individualize the quantities (precise quantification like counting).

Symbolic and non symbolic abilities develop through childhood.

3.1. Introduction

Numerical skills do not emerge de novo with language development, but are built from a biologically
determined precursor system. It is an innate ability (Ability to compare, stimate precision... are innate).

- Appears in other animal species.

- Babies can already distinguish 3 elements.
- However, symbolic abilities do depend on language.

3.2. Non-human animals

Clever Hans was claimed as a horse who could do mathematical operations (counting while stepping on the
floor). The owner would shout the operations and the horse had to answer. Some psychologists were
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suspicious and decided to study the case. They reached the conclusions that without the owner the animal
couldn't do it, so the animal couldn't count, he was understanding people's reactions.

-> In any task the expectations of the observer or the tester have a great influence in the behavior and
decision in the subject we are analysing. We need the double blind (the tested person doesn't know what's
administered, and the tester doesn't either)

If there's an innate ability, training shouldn't be needed. In these following studies they weren't trained.

1. Tamarin monkeys. Numerosity ability (quantify a magnitude, which is the number of items in a group,
realize if a group of things have more or less items than another one). Presented sequences of a
repeated sequence of items (sounds). They would then have a sequence with a different number of
items.
- Habituation paradigm (repit something so that the response of a neuron, lost of interest, etc.
changes and then do a sudden change).
- The monkey noticed the increase of the syllables used looking at the speaker, so it could
estimate.
- 4 vs 8 syllables - they could detect it
- 4 vs 6 - they could detect it
- but 6 vs 7 they couldn't
- 8 vs 12 they could
- 8 vs 10 they couldn't.
- It's not about the absolute value of the change but the proportion.

-> animals do have an innate ability on numerosity

2. Macaques. What about arithmetic operations? It’s not an habituation paradigm, but there's an
habituation phase. Game of possible and impossible additions.
- Items: screen and lemons.
- Possible: experimenter with a screen and 4 lemons inside. He shows the lemons to the
macaque. Familiarize (show him multiple times). Then, he puts 3 lemons on the shelf.
Familiarize. Test: we put the 3 lemons on the shelf, show the monkey that we add 1 more
behind the screen and then he sees 4 lemons.
- Impossible: 8 lemons. 3 on the shelf. Test: I take the 3 lemons. I show the monkey that I add
one behind the screen. When it sees that there are 8, he looks surprised.

-> Impossible additions promote stronger reactions.

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 3.3. Development in humans

(The numbers are not a sequential order)

3.3.1. Non-symbolic magnitude

Arithmetic makes two groups of elements interact (add, substract...), and numeric is to have the rough
estimation.

- Allows the perception of an approximation of the number of objects forming a group.

- Allows distinguishing between “a lot” and “a little”.
- It is an adaptive skill as it enables the automatic quantification of the environment and to act upon it
as a function of the number of present stimuli.

Habituation paradigm: The only repeated thing is the amount of elements. When you do it to babies they
lose interest. When you present any change, they are surprised and put attention again.

- Repeated presentation of a series of 3 elements (puppets, or just points), in different configurations,

produces habituation. The appearance of a series of 1 or 2 elements produces dishabituation.

Habituation in fMRI: they also use the paradigm. The disposition, size and change of the element changes
(except the number of elements). You change it now and then. Then, you compare the activation of the
brain while the habituation and the new element is showing. Result -> Activation the posterior intraparietal
cortex sulcus.

The regions that participate in realizing if something has changed are the same, and also some prefrontal
areas.

3.3.2. Symbolic magnitude in small integers

With language development comes the ability to label reality.

The representation of non-symbolic magnitude underlies the knowledge of symbolic magnitude in small
numbers.

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Mental numerical line -> all humans represent mentally the
numbers along to a mental line in different spatial positions.
There seems to be an innate preference to order things from
small to bigger from left to right. Culture will shape it.

Logarithmic -> They follow the proportions. 2 is the double of

1 so the distance is big. 2 is not the double of 3, so the space
must be smaller... natural innate non-symbolic way of measuring
quantities.

3.3.3. Symbolic magnitude in large integers

The same but with larger numbers.

3.3.4. Non-symbolic magnitude in rational numbers

Rational numbers: numbers allow comparisons.

Non-symbolic representation of magnitude in rational numbers starts at 6 months, as results from

habituation paradigm studies on ratio discrimination show.

- Test: they present groups of elements at the same time, and they have to differentiate them and tell
which one has more elements.
- Always the same ratio (you change it but keep constant the ratio). Then you change the ratio.
This way you can see that babies detect distant ratios and not close ones.

The ratio they detect depends on age.

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3.3.5. Symbolic magnitude in rational numbers

The representation of magnitude in rational

numbers is much more complex than in natural
numbers. Even in adults, performance is around
70-80% in comparison tasks with different
numerators and denominators.

Understanding that a unit can be divided in

smaller parts requires a lot of scholarization.

3.3.6. Non-symbolic arithmetic in integers

(should be put before with non-symbolic)

9 month old babies are able to detect errors in addiction and subtraction -> The same as with the monkeys
(Macaques) and the possible and impossible outcomes. Same results, they realize that the addition or
subtraction of elements is impossible.

3.3.7. Symbolic arithmetic in integers

The process usually starts around the 3 years of age with procedures based on counting, such as using
fingers to represent each number, and counting from the first (addition strategy), or recuperating from
memory the answers to simple addition problems (1 digit).

- Even in adults, more than 20% use different strategies than memory for mental calculations of 1 digit.

1 digit operations and multiplication tables as usually stored as “arithmetic facts”, i.e., in long-term memory
through repetition.

- Arithmetic facts!!-> they are real things, pieces of knowledge that work in long-term memory, and
are arithmetic since they refer to operations between different elements.

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Learning the conceptual meaning of addition helps subtraction, that of addition and subtraction helps
multiplying, and that of multiplying helps dividing. It's good to learn the concept of multiplications as a
child.

3.3.8. Arithmetic in fractions

Fractions and operations with fractions are conceptually difficult, even for adults. Most frequent errors both
in children and adults are:

3.3.9. Arithmetic in decimals

- The precision is hard even

for adults,
- Using decimals can be
really problematic for people.

4. Neural bases of numerical processing and calculus

4.1 Lesion studies

● Lesions at left-hemisphere regions related to language comprehension

○ Deficits In Comprehension And Production Of Numbers (symbolic numerical abilities)
■ Humans and other animals have a numerical sense and a magnitude estimation ability
but then with language we discretize or individualize different items because we put
an idol on them, and this is linguistic.
■ If you have problems in linguistic region, then you have problems in those abilities
(numerical comprehension and production)
● Lesions at the right hemisphere (mostly at the parietal cortex)
○ Deficits in spatial organization of quantities and in the comprehension and resolution of
abstract problems
■ Parietal cortex deals with spatial organization in vision (Where and How).
■ If there’s a lesion there, then appear deficits in organizing spatially, numerical lines,
estimating distances, or sizes, etc.
■ In order to resolve abstract problems we use a fronto parietal network. If there are
lesions there, there are problems with this.

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 ● Gerstmann syndrome
○ Lesion of Broadmann area 39 (angular gyrus) of the left hemisphere
■ Angular gyrus is related to transcription to visual objects into a language code.
○ Symptoms: dyscalculia/acalculia, dysgraphia/agraphia, finger agnosia, left-right
disorientation.
■ Dysgraphia/agraphia: inability to write
■ Dyscalculia/acalculia: inability to write numbers and inability to operate with numbers.
■ Finger agnosia: confuse which finger is which

4.2 Neuroimaging studies

Key regions in the network underlying numerical processing and calculus are:

❖ Intraparietal Sulcus (IPS):

➢ Key region for the numerical sense, magnitude representation in abstract terms. Ability even
macacs and monkeys show
➢ Lesions here appear to cause problems in numerical attitudes (symbolic and not symbolic).
➢ Abstract representation of numerical magnitude
❖ Posterior Superior Parietal Lobule:
➢ Attention & Multi-digit operations
➢ We situate in space multi-digit operations and we get the solution.
❖ Angular gyrus:
➢ Language dependent processes (multiplication tables, simple operations, “arithmetic facts”)
➢ Arithmetic facts: when you don’t need to perform an operation. Ex: 12x12=144. (las típicas
que te sabes de memoria y que no tienes que pensar)
❖ Fusiform gyrus:
➢ Arabic number processing
➢ Ability to make aloud a meaning of a symbol, perceived as a whole. The symbol 6 there’s no
reference to how it sounds, and that’s why we used it internationally in all the countries that
adopt arabic number processing (six, seis, sis, etc.)
➢ You just need a phonological look.

Parietal Lobe

➔ Intraparietal Sulcus (IPS)

◆ Neural substrate for number sense, magnitude estimation.
● Ex: There’s more elements in one group than another
◆ Bilateral Activation Magnitude Estimation Tasks

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 ● People discuss that can be divided in 2 different domains. One is a non-symbolic
approximation number system (ANS) and another is symbolic in which we individualize
items, discritizises, such few (even babies can individualize 2 or 3 elements).

◆ Habituation paradigm in fMRI. Repeated magnitudes diminish IPS activity, different

magnitudes enhance it.
● They do a study that consists of doing an Habituation paradigm in which they change
numbers and shapes, etc.
◆ Processing symbolic magnitudes (digits) and non-symbolic magnitudes (points, angles…) in
different formats (size, time, speed, angle...)
◆ Task: Comparing magnitudes
● Numbers (symbolic)
○ So they try to individualize the activities related only to a change in number. If
the habituation is 16 stimuli (images) that change in organisation, in size, etc.
and comparing these changes but only to the number change the intraparietal
sulcus is more active.
● Angles (non-symbolic)
○ Even we can get it with other non-symbolic magnitudes (angles)
○ The activation of the intraparietal sulcus is common
● Lines (non-symbolic)
○ And estimate the length of a line.
○ The activation of the intraparietal sulcus is common
◆ Results:
● Bilateral activation of IPS in all cases
➔ Angular gyrus
◆ Involved In Language,visuo- verbal coding and verbal memory processes.
◆ Belongs To The Linguistic System for calculus.
◆ Encodes “arithmetic facts” (automatic operations stored in verbal memory): multiplication
tables, 1-digit additions.
● The angular gyrus is mostly involve in arithmetic facts
◆ Aids In Comprehension And expression of numbers in verbal format (symbolic).
◆ Active in known arithmetic operations.
◆ Study→ Task
● Training in multiplications
○ They train subjects in multiplication
● fMRI: Comparison of active areas while performing new multiplications vs. those
trained
○ Subjects had to do a figural-spatial task in which they had to count the angles
that they had.
○ They put the subject multiplication that they already knew the answer and
other that they don’t train.
○ Comparison between untrain brain and train brain.
◆ Results
● Untrained multiplications activated the IPS and the prefrontal cortex
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 ○ Comparing magnitudes, attention and working memory are required.
◆ When you don’t know the answer of a multiplication and you have to
compute it you see activity in the parietal sulcus, prefrontal cortex,
superior parietal lobule.
◆ But once you know the answer the angular gyrus is activated (more
activation on the right side than on the left)
● Trained multiplications activated the angular gyrus
○ Verbal memory.
➔ Posterior Superior Parietal Lobule
◆ Multimodal area above IPS.
◆ Attentional,visuospatial and spatial working memory tasks involved in calculus.

Frontal Lobe

➔ Prefrontal cortex
◆ Involved In Executive System functions. The more complex the task, the more active.
● Numerical Sense
○ Ordering numbers in the mental line
● Arithmetic facts
○ Retrieving, ordering and using them (fronto-parietal networks including
prefrontal-anterior cingulate connections).
● Logic and reasoning
○ Planning, working memory and attention in problem solving (Pesenti et al.,
2001).
● Error correction
○ Monitoring results and correcting errors (Menon et al., 2002).

Temporal Lobe

➔ Fusiform gyrus
◆ Processing of Arabic numbers (Right Hemisphere dominant)
● doesn’t hold all the information about how it sounds (és lo que hem dit abans)
◆ Processing of faces (Right Hemisphere dominant) and the visual shape of words (Global
reading; Left Hemisphere dominant).

4.3 Numerical neurons

● IPS (intraparietal sulcus) and mPFC (medial prefrontal cortex) contain multimodal “Numerical
neurons” that respond selectively to a particular quantity (around 20- 30% in mPFC and 15-20% in
IPS).
○ In these experiments, they presented the monkeys with a delayed match-to-numerosity task.
○ A stimulus is presented which contains a numerous of items (sizes change). They will be
presented with another sample that matches with the sample or not. So, the monkey has to
be balanced, it has to think if it matches or not.
○ The experimenters thought that their neurons preferes a specific number of items.

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 ○ Els puntets son Potencials d’Acció

○ This experiment contain other conditions:

■ Sequential visual number task
■ Sequential auditory number task
● they did it with sounds
■ Supra-model number-turning curves
■ Number network in the monkey brain
■ Number network in the human brain
○ The experimenters thought that these neurons in humans will be more related to medial
prefrontal cortex, and intraparietal sulcus response is rewardless in the modality of the visual
or auditory modality preferring one quantity.

5. Dyscalculia
● In a typically developing child, the critical period for numerical learning ranges from 5 to 7 yrs old.
○ Kids could start studying math later at school!
● Some mathematicians think we should completely change the way we teach math.
○ We spend too much time teaching calculus,when that can be
done by a computer.
● Is math difficult?
○ If you see the performance of a child in math exercise you will see the campana de Gauss.
○ At least one or two people in class will have this disability and they get very frustrated. We
have to be aware of this potential difficulty.
■ Maybe they can plan a reason properly and they could
theoretically solve a math problem.

● Difficulties In math can have Primary origin or a secondary one following a lesion.
● Acalculia is the loss of ability to perform tasks related to numerical processing and/or calculus as a
consequence of some neural pathology.
● Developmental Dyscalculia Specific disorder affecting the correct acquisition of numerical abilities
without apparent cause. It interferes with academic performance and daily live numerical activities
(ex. Street Numbers, phonenumbers...) and cannot be explained by any sensory, neurological or
mental deficit (Shalev, 2004). It is often associated with dyslexia and ADHD (25-30% comorbidity).

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 UNIT 10 - EXECUTIVE SYSTEM
1. Introduction

1.1 Executive functions in our daily lives

Plan grocery shopping and cooking dinner for friends:

1. Select a recipe.
2. Write a grocery shopping list.
3. Plan the path to visit all stores in the shortest time possible.
4. Do not get distracted looking at showcases or talking to neighbors
5. Read the recipe and prepare and cook all ingredients in order.
6. Get the house ready.

Most people do not find this sequence particularly difficult, but people with executive dysfunction can’t do
it.

Planning is an executive function. It's necessary memory sometimes, planning is using our resources to do
a mental map to do what we want to do. Many people with problems in self-control have troubles with
keeping their goal.

1.2 Concept of executive function

Executive functions are skills rendering a person able to participate successfully in an independent, goal-
directed behavior.

➔ Executive functions control cognitive processes so that appropriate actions according to

spatiotemporal contexts are selected.
➔ Executive functions entail:
➔ Formulating Objectives.
➔ Planning the consecution of those objectives.
➔ Successfully executing those plans.

1.3 Modulating factors of executive functions

❖ Internalized information
➢ Dorsolateral Prefrontal Cortex
■ Registry of recent events in temporal order
■ Can be related to objects or actions
■ Related to working memory
● Important in decision making.
❖ External information
➢ Dorsolateral Prefrontal Cortex
■ Inhibition of external signs to focus on own objectives
● When there are many people around you and someone is talking to you, you
have to focus on that person and ignore the rest.

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 ➢ Orbitofrontal Prefrontal Cortex
■ Associative learning between stimuli and reward
❖ Context
➢ Orbitofrontal Prefrontal Cortex
■ Social role (parent, offspring, sibling, lover, friend....)
● it’s different to cook for your parents, lover, close friend, etc.
● Depending on your social role you will modulate your behaviour.
■ Behaviour adaptation to social environment (library, club…)
❖ Self-knowledge
➢ Orbitofrontal Prefrontal Cortex
■ About our personal experience and life goals

1.4 Executive functions

➢ Organization and planning

➢ Anticipation
➢ Goal selection
➢ Sustained attention
➢ Problem solving
➢ Working memory
➢ Cognitive Flexibility
➢ Inhibitory Control

1.5. Function of executive functions

Executive functions supervise and regulate the activity of other cognitive processes in a flexible and goal-
directed manner.

In executive function we are dealing with goal-oriented behavior, you have a purpose (external factors) to
behave, that’s different from motivation (internal factors). I mean, we are dealing with goals, we have a
purpose to get there, and then we need to organize all our behavior and factors that could influence in
reaching the goal, so control them.

The key region is the prefrontal cortex. It would redirect the groups of information between other regions,
so redirects the functional connectivity.

So to speak, they are a general organizer of brain function, such as a railroad switch operator.

1.6. Alteration of executive functions

Alteration of executive function impacts virtually any cognitive and emotional function: personality disorders,
affective disorders, mood disorders (ex: phineas gage), character disorders, motivation disorders, attention
disorders, language disorders, memory disorders, problem solving disorders, reasoning disorders and
thought disorders. EX concrets: dementia (they become cascarrabias and this is something can happen
because prefrontal degeneration, they can lose impulse control and become a viejo verde...).

It could be because there is a lesion or an imbalance in neurotransmitters.

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 2. Development of executive functions

2.1. Phylogenetic perspective

From a phylogenetical point of view (evolutionary perspective) the

prefrontal cortex is one of the regions that developes late. The prefrontal
cortex is amongst the last regions to develop phylogenetically.

It shows an accelerated growth in primate species. Prefrontal cortex is so much larger in humans than in
other species. Ex: cat → 3,5% of total brian volume, dog→ 7%, chimp→ 17% and human → 30%. Are we
better than cats? If we are going to extinguish ourselves by destroying our world, we have not been that
successful evolutionary talking. But what we can not deny is that this ability to create these virtual walls in
which we use language to perceive what is going to happen and adapt our behavior extremely fast is
extraordinary. Because, in fact, a change in behavior of other species takes years and years of evolution,
and by culture we can change a behavior in only 1 generation. We adapt incredibly fast.

2.2. Ontogenetic perspective

The prefrontal cortex is amongst the last regions to mature. First, we develop primary areas, which are
essential in babies. Then we develop primary areas of association, ones which associate the same kind of
information. And the last areas to develop are the secondary areas of association, because it takes different
types of inputs.

Structural maturation ends at 16-18 years old. By maturation we say that reach the adult-like point. Using
drugs before this time, it really impacts brain functioning.

Prefrontal cortex maturation (as measured by the time course for synaptogenesis) starts before birth and
lasts until late teenage years. We have the peak of synapsis at 3 years and it stills until 15 years, more or
less.

The primary regions have a large number of synapses when we are born until 2/3 years when we have lost
most of them (we are perfectly mature in these regions).

Executive functions also are the last cognitive functions to mature. There is a clear relation between
maturation and executive function development.

Prefrontal cortex maturation (as measured by the time course for myelination-integrity of withe substance)
starts before birth and lasts until late teenage years.

DTI longitudinal study from 5 to 21 years old shows how not all white matter tracts mature simultaneously,
and that those connecting the prefrontal cortex are the last ones to mature.

During teenage years, a maduration boost occurs for cortico-cortical and cortico-subcortical fibers,
especially involving prefrontal cortex connections, underlying executive functions. That’s the reason why
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society says something is illegal or legal because of being an adult, like drinking, driving, smoking… because
executive function until those years is not fully developed.

2.3. Ontogenetic perspective - functional:

● Executive functions start developing during the first years of life and continue to do so all life.
● Some periods exhibit greater changes, such as the teenage years.
● Different development speeds occur naturally between functions and individuals (great intrinsic
variability).
● The development of executive functions is tightly linked to the structural maturation of the prefrontal
cortex and its networks.
● Academic skills, motor learning and social skills require great executive function capacities.
● The more complex the task, the greater the involvement of executive functions.

In executive function we move from an externally controlled behavior, because behavior needs structuring,
because being the main purpose of executive function to guide a goal directed behavior, so to have a
purpose to act.

Executive functions
are developed from
childhood, under
adult influence, until
they become
independent.

3. Anatomy and function of the executive system

Executive functions rely on the activity of the prefrontal cortex (PFC; associative cortex) and its connections
with other brain areas forming networks.

The PFC is delimited according to:

● Cytoarchitecture: thick layer IV.

● Connectivity: receives afferences from the dorsomedial thalamus.

The PFC can be divided in:

1. Dorsolateral prefrontal cortex (dlPFC).

2. Orbital prefrontal cortex (oPFC).
3. Medial (Paralimbic) prefrontal cortex (mPFC).

3.1. Connectivity of the prefrontal cortex

The OFC is widely connected to cortical and subcortical structures:

● Cortico-cortical connections
○ Amongst PFC regions
○ With other associative areas (parietal and temporal)
○ With the limbic system (through mPFC and oPFC), in the sense of making a decision or related
to motivation, ex: i still love him? or should i keep doing this?

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 ● Cortico-subcortical connections
○ With the basal ganglia and the thalamus (prefrontal-basal
ganglia-thalamic-prefrontal loops)
■ There are 5 fronto-subcortical loops:
● Motor
● Oculomotor
● Dorsolateral Prefrontal
● Lateral Orbitofrontal
● Anterior Cingulate
■ Loss of integrity in these circuits is associated to
executive dysfunction, observed in pathologies
such as:
● ADHD
● Schizophrenia
● White matter lesions due to small vessel
vascular pathology.
○ With the hypothalamus (mPFC and oPFC)
○ With the amygdala, ex: the teacher says that tomorrow there will be a surprise exam, so we
get anxious because we plan and think how we can study all the topics in just 1 day. And then
you get stressed, so you feel stressed → hypothalamus, autonomus nervous system.
○ Esquema: diferent cortical regions (prefrontal regions) → that send information to the
striatum (it integrates this information) → it goes to the globus pallidum with substantia nigra
(parts of the basal ganglia that output information to the thalamus, and thalamus bring it
again to the cortex).

3.2. Subdivisions of the prefrontal cortex (this is so important!!)

● Dorsolateral Prefrontal Cortex (dlPFC): cognitive aspect of executive function: decision making,
reasoning, planning, solving problems, directing attention, evaluating the consecution of a goal.
○ The connections with the parietal cortex are important in working memory.
■ Ability to maintain actively (and temporally) relevant information to complete a task.
For instance, remembering the data of an arithmetic problem or the direction to which
we are delivering a package.
○ dlPFC functions are chiefly explained by its connections with the parietal cortex.
● Orbitofrontal cortex (oPFC): is much more related to emotion regulation and self-regulation
(remember social behavior, is not an executive function, but we adapt our behavior by the social
context). This impacts your decisions, because it is connected through the loop that we explained
just before. EX: if you would not feel bad, anything would care (like a psychopath).
○ Connectivity:
■ Receives information from all sensory modalities. Region important in putting
together what we see or evaluate with our feelings.
■ Influences the autonomous nervous system (respiration, blood pressure...) and the
endocrine system (emotional responses). Amygdala & hypothalamus.
○ oPFC functions:
■ Adapting behavior to context: oPFC deals with life in society and hierarchy. People
with lesions in this region know social scripts (rules) but do not apply them.

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 ■ Behavior flexibility: lesions at oPFC cause persistent behavioral patterns,
perseverations, loss of ability to change tactics after failure.
■ Associative learning: encodes reward and punishment values of primary (ex: food) and
associates them to secondary stimuli, so that they participate in motivational
behaviors such as feeding, emotional and social behaviors.
■ Encoding information in declarative memory
■ Emotional and motivational processes
● Medial Prefrontal Cortex (mPFC; aka. Paralimbic): more related to the motivational aspect. Actually
lesions in the anterior cingulate can evoke feelings of apathy.
○ mPFC connectivity shows an intermediate position between the limbic system and the
neocortex.
○ mPFC is considered part of the limbic and the Prefrontal systems.
○ Functions:
■ Motivation and behavior initiation.
■ Connection between volitional, motor, cognitive, emotional and memory processes.
■ Regulation of autonomic functions.
■ Selective attention

3.3. Neurotransmitters involved in executive function

● Acetylcholine
● Histamine
● Serotonin
● Noradrenaline
● Dopamine

4. An example: Attention

4.1. Definition

“Everyone knows what attention is. It is taking possession of the mind, in clear and vivid form, of one out of
what seems several simultaneously possible objects or trains of thought. (Selectivity: inhibit distractors to
have maintained attention) Focalization, concentration of consciousness are of its essence. It implies a
withdrawal from some things in order to deal effectively with others, and is a condition which has a real
opposite in the confused, dazed, scattered brain.”

- James, W. (1890), father of american school of psychology.

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William James already realized that attention can be effortful, implies perception and acts as a filter on the
outside world.

According to Breedlove & Watson, 2017 (Behavioral Neuroscience, 8th Ed.):

- Attention, also called selective attention, is a state or condition of selective awareness or perceptual
receptivity, by which specific stimuli are selected for enhanced processing.

4.1.1. Selectivity

Being in a state of enhanced alertness and to focus on some information (objects thought, etc.) and not on
others are different things.

- Selectivity is the key attribute, as opposed to:

- Arousal, a global, nonselective level of alertness of an individual.

4.2. Classic neuroscience of attentional networks

A fundamental distinction in the study of attention is between the control of attention (directing the
attentional beam) and the implementation of attention (illuminating objects by the beam) - Posner &
Petersen (1990)

❖ Control of attention -> deals more with the direction of the spotlight, the most executive part of
attention (where do I want to attend? / maybe an external stimuli shifts our attention -> this would
be the control of attention, the shifting and engaging). All the resources go to this biased target.
➢ Controllers
➢ Ex: If we’re looking for our phone, circular objects won’t be biased, anything made out of
wood can’t be enhanced either… the processing of those features (those of a phone) will be
biased.
❖ Implementation of attention -> the fact that when the spotlight is on the object and we have a train
of thought more clear and that we follow (while other trains of thought that we may have we are not
following) is the implementation of attention.
➢ Processors
➢ Where attention is implemented
➢ The processing in itself
➢ Ex: I need to find my phone so my controller grabs all the processors that it needs to find the
phone like the visual one (and I start doing a visual search) and the controller will guide it.
The controller will have a saliency map which is what is biased so that the features of being
rectangular, having the color of our phone, etc. will be biased. The fact that the eye moves
to look for it means that we’re processing data in that location and not in others (visual bias).
You can’t find it so you ask someone to call you. Then, the controller grabs your hearing and
makes you pay attention with it, you are processing more the controller.

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The construct of attention has classically been operativized using Posner’s Attentional Networks Test (ANT)
(controllers!)

➢ There are 3 tasks that helped them to try to study different networks and brain functions related to
different aspects of the control of attention.
○ Alerting = A momentraily increase of arousal (which is not selective)
■ Ex: Someone is falling asleep and you suddenly shout. That stimulus made them alert,
but they’re alert to whatever.
■ Task: A cross in the middle of the screen and you have to look at it. Then, there’s a
cue (like a sound that doesn’t tell you where the number will appear) and there will
appear a number. You will have to press a button if the number is greater than 5 and
press another one if it’s smaller.
■ More activation of the Locus Coeruleus (related to the state of arousal and dilatation
of the pupils); Right frontal and parietal cortex. The Nt that modulates the network is
the Norepinephrine.
○ Orienting = An instruction tells you to direct the focus of attention to a particular
location/object/feature or whatever (it is selective).
■ Task: The cue is an arrow pointing where the number will appear. You can bias your
attention towards that location. If the cue predicts correctly the appearance of the

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 stimuli, the processing is much better. If not, you’re worse than in the alerting task,
which means that there’s a cost in orienting your attention if the stimulus of relevance
appears outside of the focus of attention.
■ Superior Parietal Lobule (we saw it in the number opic and the visuospatial one),
Temporoparietal Junction (Theory of mind), Frontal eye field (spatial behavior),
superior colliculi (vision). The NT that modulates this network is the Acetylcholine.
○ Conflict = You have several things that reclaim the oriental attention but you need to focus
only on one, or, what happens when there’s an association between stimulus and a response
but at some point you have to inhibit it (examples of conflicting situations)
■ Attention will have another aspect dealing with the conflicting information
■ Task: Tell the color of the word, not read the word. We’re obviously faster in the
congruent one than in the incongruent because there’s a conflict of information, we
have to inhibit the initial response that we would do. It could also be that you have to
say the letter of the middle (A/B) or that you have to press one button if the arrow
points up or down (if we press with the right, we’re faster with the arrow pointing
upwards).
■ Anterior cingulate cortex (related to conflicting information, effort that we have while
dealing with it, connections with the limbic system - emotionality takes part in the
conflict), the lateral and ventral part of the prefrontal cortex, basal ganglia. Dopamine
is the one signaling.

4.2.1. Orienting

“The orienting network is focused on the ability to prioritize sensory input by selecting a modality or
location.” (Petersen and Posner, 2012).

Following the spotlight metaphor (the spotlight moves and enhances the processing of something), the
location of the focus of attention can be driven by:

- Exogenous attention: External stimuli (due to a stimulus saliency or valence) cause the shift.
- It’s because of probability/statistics.
- Ex: If there’s a picture in black and whithe and there’s suddenly a red spot, it catches
our attention since the probability of finding color in the picture was close to none. If

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 our teacher suddenly increases the pitch of his voice, it will also catch our attention
since we know that it doesn't sound like that and we weren’t expecting it, again,
because of probability. It’s unlikely, so it make it salient.

- Endogenous attention: Internal goals (due to a stimulus relevance to the task) cause the shift.
- Ex: I’m hungry, which means that anything that’s food related will grab our attention. Our
internal goal tells us that it’s relevant. It’s different than moving the focus of attention
voluntarily.

Dorsal Fronto-Parietal Network (Top - Down)

Highly active when we voluntarily pay attention to something. I decide what to look at and what I want to
process better. We enhance the processing of certain features.

❖ IPs: IntraParietal sulcus

➢ Some cells behave as if preparing for an attentional shift (they selectively increase firing to a
covertly attended target).
■ Coverty = atenció encoberta de percepció
■ The cells increase their firing toward locations of the targets when we covertly attend
to them without moving the eyes.
➢ Neurons in the LIP (Lateral IntraParietal sulcus, monkey homologue of human IPs) encode a
saliency map (priority map) that controls the direction of attention and eye movement
planning between important stimuli (regardless of perceptual modality).
■ These neurons encode for the number of items (from another topic) and are very
active in a network when you are paying attention to something.
➢ Interrupting/lesioning IPs (intraparietal sulcus) function affects voluntary shifts of attention.
❖ FEF: Frontal Eye Fields
➢ Important for establishing gaze (eye movement) according to cognitive goals (top-down)
➢ Biases neural activity in visual cortex (tunes neurons to process selected stimuli)
■ Ex: looking for Wally is voluntary, but when it pops out in our vision field, it’s due to
its aliency.

Ventral Fronto-Parietal Network

When the thing that attracts your attention is external. When we are engaged in something we have to be
able to disengage in case we have to pay attention to something else (we’re engaged in studying but we
sense a burnt smell, we have to be able to engage in this and disengage from studying -> we go back to
the top-down to engage in that something else).

❖ TPJ: Temporo Parietal Junction

➢ Role in shifting attention to a new location after target onset, especially if unexpected.
■ Break one engagement to get into another
➢ People with damage at TPJ respond poorly to unanticipated targets.
➢ TPJ could act as a circuit breaker, overriding the current attentional priority to redirect itto
something new.
➢ Strongly right lateralized system
■ (congruent with a syndrome, hemispatial neglect - percepció).

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 ❖ VFC: Ventral Frontal Cortex
➢ With the inferior frontal gyrus and the medium frontal gyrus
➢ Involved in working memory
➢ May provide crucial information about novelty by comparing present with past stimuli.
■ “This isn’t what I was expecting”

Hemispatial neglect:

The lesions coincide with the Ventral Fronto-Parietal regions. What happens to these people is not that they
don’t see, but everyhting that is on their left hemiiels of vision doesn’t pop out, it doesn’t call for their
attention so they are neglected. They are anosognosic as well, since they are not aware that they have this
disorder.

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4.3. Effects of attention

“Immediate effects of attention are to make us perceive … better than otherwise we could” (James W,1890).

These are the effects that attention has:

- Facilitation
- Higher Accuracy
- Faster Reaction Times

BOLD measures changes in the levels of oxygen in our blood (the more the harder it’s working). When we
tell someone to h¡¡shift their attention in something, we can see an enhancement on the flow of oxygen.

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When the stimulus appears there’s more activity, but, when we’re paying attention, there’s an increase as
well.

(Look at the graph of the Visual Cortex and the FEF)

When attention is directed towards one ear it shows enhanced electrical activity.

Graph 1: When the animal pays attention, neurons are active to stimuli even in low contrast than when they
are not attending (actually, with no attention there’s 0 reaction to low contrast). This means that we see
better when we pay attention since teh neurons are firing more.

Graph 2: When the animal pays attention to a particular frequency, even if the frequency is the one it likes,
firing is increased. If that frequency is a frequency the neuron doesn’t like, there’s a decrease. Neurons react
more to what they like and less to what they dislike.

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Graph 3: Attention synchronizes action potentials.

4.4. A plausible mechanism for selective attention

The timing between spikes is a really important code in the brain for communication, and the spikes get
more synchronized in local populations when we’re attending. All the neurons work together for the same
goal.

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