Neuroni

specchio e linguaggio

Gloria Gagliardi
gloria.gagliardi2@unibo.it

 SOMMARIO
Lezione del 17/11/2014:
Quadro generale

Cenni di morfologia delle aree cerebrali

Le aree motorie (nella scimmia e nelluomo)
Neuroni canonici e aordances
Neuroni specchio

Neuroni specchio, leMura delle intenzioni ed empaNa

Embodied cogni2on e teoria della mente
 SOMMARIO
Lezione del 19/11/2014:
Perch la scoperta dei neuroni specchio dovrebbe interessare al
linguista? In che modo inuenza la teoria del linguaggio?

Filogenesi del linguaggio

Ontogenesi del linguaggio
Azione/Gesto/Linguaggio
(studi di V. Volterra e ISTC-CNR)

Case study acquisizione aNpica: lauNsmo

(V. Gallese; L. Brandi e S. Lucchesini)
Neuroni specchio, percezione e comprensione linguisNca
Liv. Fonologico (Motor theory of speech percep2on)

Liv. Lessicale-semanNco
MORFOLOGIA delle AREE CEREBRALI
(Rizzola_ e Sinigaglia, 2006)

CitoarchiteMura del cervello di scimmia (Cercopiteco), mappa di Brodmann

MORFOLOGIA delle AREE CEREBRALI

CitoarchiteMura del cervello umano, mappa di Brodmann

 AREE MOTORIE
Aree 4 e 6 di Brodmann,
corteccia agranulare

Area 4, corteccia motoria
primaria, F1
Area 6, disNnta in 3 regioni
principali (ciascuna divisa a
sua volta in una parte
rostrale ed una caudale):
Regione Mesiale: F3, F6
Regione Dorsale: F2, F7
Regione Ventrale: F4, F5
 AREE MOTORIE
Area 4
Controllo del movimento volontario
Organizzazione somatotopica

Semiunculus di C. Woolsey Homunculus motorio di W. Peneld

 Controllo del Movimento
La mano ha una struMura
anatomica molto complessa:
39 muscoli (estrinseci ed
intrinseci) che agiscono su
18 giunN
23 gradi di libert dal
punto di vista cinemaNco

COME VIENE CONTROLLATO

IL MOVIMENTO?
 Area F5 (scimmia)
PROPRIETA GENERALE DELLAREA F5 (Rizzola_ et al. 1988)
Neuroni dellarea F5 hanno propriet pi complesse di quelle
della corteccia motoria primaria. Codicano qualcosa di pi
astraMo del singolo movimento: scaricano in concomitanza di
a_ motori specicamente goal-related.

The funcNonal properNes of neurons located in the rostral part of inferior
area 6 were studied in awake, parNally restrained macaque monkeys. The
most interesNng property of these neurons was that their ring correlated
with specic goal-related motor acts rather than with single movements
made by the animal.

All together these neurons form a vocabulary where proximal and distal
movement necessary for reaching, grasping, holding and bringing the food to
the mouth are represented.
 Una dimostrazione
Umilt et al. 2008
Primates are able to interact with objects not only by using their natural
eectors, but also by using tools. Common tools, such as sNcks, stones, and
rakes, act basically as funcNonal extensions of natural eectors. With
pracNce, they become parts of the agents body schema.

LO STUDIO:
Two adult macaque monkeys
(Macaca nemestrina) were used.
Single-unit acNvity was recorded
from areas F5 and F1
Before recording, the monkeys
were trained to grasp food placed
in front of them by using two
types of tools.
To grasp the object with normal
pliers, the monkey has to close its
hand (A), whereas with the
reverse pliers, the monkey has to
open its hand (B).
What could be the mechanism that allows a transformaNon of a goal into
appropriate movements even when an opposite sequence of movements is
necessary to achieve the goal? Our ndings show that, aqer learning, the
correct movement selecNon occurred immediately as soon as the monkey
grasped one or the other type of pliers.

goal-centered organizaNon of primate motor cortex


 Neuroni canonici
Murata et al. 1997

Visual and motor properNes of single neurons of monkey ventral premotor cortex
(area F5) were studied
The employed objects were six dierent three-dimensional (3-D) geometric solids.

FINDINGS

Two main types of neurons were disNnguished: motor neurons and visuomotor
neurons.
Motor neurons discharged in associaNon with grasping movements. Most of them discharged
selecNvely during a parNcular type of grip. Dierent objects, if grasped in similar way,
determined similar neuronal motor responses.
Visuomotor neurons also discharged during acNve movements, but, in addiNon, they red also
in response to the presentaNon of 3-D objects.

every Nme an object is presented, its visual features are automaNcally (regardless
of any intenNon to move) translated into a potenNal motor acNon. This potenNal
acNon describes the pragmaNc physical properNes of the objects.
 Neuroni canonici
CaraMerisNche:

Neuroni visuomotori che scaricano sia durante gli a_ motori, sia
alla presentazione di ogge_ 3D.

Ogni volta che un oggeMo viene percepito, le sue caraMerisNche

siche vengono automaNcamente tradoMe in un potenziale
motorio di azione.

La visione di un oggeMo a_va immediatamente la selezione

delle propriet siche che permeMono di interagire con esso
AFFORDANCES


 Aordance
ConceMo introdoMo da Gibson
The aordances of the environment are what it oers the animal, what it provides or
furnishes, either for good or ill. The verb to aord is found in the dicNonary, the noun
aordance is not. I have made it up. I mean by it something that refers to both the
environment and the animal in a way that no exisNng term does. It implies the
complementarity of the animal and the environment.
(Gibson 1979, 127)


DeFelice (2013)
L'autore denisce le aordances come possibilit d'azione oerte dagli ogge_: il
conceMo non va ridoMo per alle semplici propriet siche e visivamente percepibili
degli ogge_ in s, ma intrinsecamente relazionale, in quanto implica che sussista
una complementariet tra gli agenN e l'ambiente (e gli ogge_ che ne fanno parte)

DeFelice (2014), Denizione di Aordance:
invariant set of objects physical properNes that oer possibiliNes for acNons.

 Aordances e neuroni canonici
Where are Aordances grounded? CANONICAL
NEURONS CIRCUITS

La percezione di propriet di ogge_ manipolabili

a_va una sorta di simulazione dellazione nei
circuiN cerebrali

Stre_ssima relazione tra azione e percezione:

labilit di agire dipende dalla abilit di percepire.
 Neuroni Canonici nelluomo
Grezes et al. 2003a

S2muli: color video recordings (3.5 s each) of objects, grasping
pantomimes, and objects being grasped.

The subjects viewed the videos and were required either to passively
observe or to execute the appropriate grip on a manipulandum.
In the observaNon context, subjects observed an object (OO), observed a
grasp (OG), or observed an object being grasped (OGO).
In the execuNon condiNons, the subjects executed the grasp appropriate for
the object that they viewed (EO), imitated the pantomime they viewed (EG),
or imitated the hand grasping an object (EGO).

Two dierent objects were used; one had large opposiNon axes and would
normally be grasped with a power grip. The other object was small and
would normally be grasped by a precision grip.
 Power grip Precision grip
(thumb + surface of the palm + all other ngers) (thumb + index nger)
 fMRI scanning

The manipulandum employed for responses

had two components. The rst component
consisted of a wooden cylinder [], that was
used for the power grip response. AMached to
the top part of the cylinder, the second
component was made of a 2-cm-square and
1.5-cm-thick piece of wood and was used for
the precision grip response. Both components
contained a force transducer to record grip
responses.

Comparing those data with electrophysiological ndings in monkeys, the concomitant

acNvaNon of parietal and pre-motor areas during passive observaNon of an object and
during execuNon of a grasp toward that object seems to correspond to the circuit
shown in the macaque brain, that is, the parietal area AIP and the premotor area F5,
where canonical neurons have been recorded.
Buccino et al. 2009
TMS: Transcranial MagneNc SNmulaNon1
STIMULI: The sNmuli set was made of 6 2 2 sNmuli, each object being
represented in two horizontal orientaNons (one compaNble with a right-hand
grasp, the other with a leq-hand grasp) and according to two dierent types of
handle (one normal and compaNble with a hand grasp, the other broken).

Motor programs for grasping objects are specically related to some specic
pragmaNc features of objects like, for example, handle orientaNon.
The recruited motor programs are ne tuned to privileged components so that as
soon as these are violated, the related motor programs are consequently
interrupted, as tesNed by MEPs recorded when objects were presented with the
broken handle on the right side

1 Applicazione di un impulso magneNco alla corteccia. Se limpulso applicato alla corteccia motoria possibile registrare i potenziali evocaN
motori (MEP) nei muscoli controlaterali mediante eleMromiograa.

 Neuroni Specchio
ScoperN negli anni 90, a Parma [Rizzola_, Gallese, Fadiga, Fogassi]
Primi arNcoli: Rizzola_ et al. 1996; Gallese et al. 1996

Dove? Area F5 della scimmia

Cosa sono? sistema di neuroni parNcolari, a_vi sia quando il soggeMo compie lazione,
che quando lazione compiuta da un altro agente e semplicemente osservata.

cfr. Gallese et al. 1996:
Registrazione di 532 neuroni dellarea F5.
92 hanno propriet mirror.
Di quesN:
51 si a_vano durante losservazione di una singola, specica azione
compita con la mano.
38 si a_vano per 2 o 3 azioni.
3 si a_vano quando la scimmia osserva il ricercatore prendere cibo
con la bocca o con la mano

25: mirror-like, rispondono allosservazione di azioni della mano ma non
hanno propriet motorie.

 Neuroni specchio nella scimmia:
funzione e caraMerisNche
HP: il sistema specchio gioca un ruolo nella comprensione delle azioni

Il traMo visivo che a_va i neuroni specchio linterazione tra agente e
oggeMo: la vista del solo agente o del solo oggeMo non produce scarica.

I neuroni specchio della scimmia non scaricano se lazione viene mimata
dallagente.

I neuroni specchio si a_vano quando lanimale pu prevedere la nalit

dellazione
studi molto noN:
Umilt et al. 2001
Kohler et al. 2002; Keysers et al. 2003



Umilt et al. 2001

Two basic experimental
condiNons:
full vision condiNon
[the monkey was shown a fully
visible acNon directed toward an
object]
hidden condiNon
[the same acNon was presented
but with its nal criNcal part (hand-
object interacNon) hidden].

The results showed that the

majority of mirror neurons
responded also in the hidden
condiNon.
 [] a subset of mirror neurons becomes acNve during
acNon presentaNon and also when the nal part of the
acNon, crucial in triggering the response in full vision, is
hidden and can therefore only be inferred.

This implies that the motor representaNon of an acNon
performed by others can be internally generated in the
observers premotor cortex, even when a visual descripNon
of the acNon is lacking. The present ndings support the
hypothesis that mirror neuron acNvaNon could be at the
basis of acNon recogniNon.

Full visual informaNon about an acNon is not necessary to

recognize its goal. AcNon understanding could be based on
a mechanism that can trigger the internal motor
representaNon of the acNon.
 Audiovisual mirror neurons
Kohler et al. 2002/Keysers et al. 2003

STIMULI= noisy acNons
Peanut breaking
Paper ripping
PlasNc crumpling
Metal hi_ng metal
Paper shaking
SNck dropping

three sensory condiNons:

vision-and-sound (V+S)
vision-only (V)
sound-only (S)
 [] some neurons in the ventral premotor
cortex (area F5) of the monkey responding
during the execuNon of acNons also respond to
the vision and/or the sound of these acNons.

[] for half of the tested audiovisual mirror

neurons, the amplitude of the response does
not dier signicantly whether the preferred
acNon is heard, seen or both heard and seen.

This nding is important, as it suggests that the

neurons code the acNon in an abstract way,
which does not depend on the source of
informaNon (auditory or visual) from which the
evidence about the presence of the acNon is
taken.

cornerstone in the evoluNon of language?

 Non solo F5: Mirror-neuron circuit
(Rizzola_ e Sinigaglia, 2006)
Il sistema dei neuroni specchio bilaterale e include ampie porzioni della
corteccia parietale e premotoria.

F5

Area PF of Von Economo

(rostral part of the inferior
parietal lobule, IPL)
It receives input from STS and sends an
important output to the ventral premotor
cortex including area F5. PF neurons are
funcNonally heterogeneous. Most of them
(about 90%) respond to sensory sNmuli, but
about 50% of them also have motor properNes
discharging when the monkey performs specic
movements or acNons).

STS (superior temporal sulcus)
STS neurons respond to the observaNon of
acNons done by but do not appear to be
endowed with motor properNes.)

The corNcal mirror neuron circuit is formed by two main regions: the
rostral part of the inferior parietal lobule and the ventral premotor
cortex. STS is strictly related to it but, lacking motor properNes, cannot
be considered part of it.
(Rizzola_ & Craighero, 2004)


 Neuroni specchio: funzione
The mirror system provides a way to match observa2on and execu2on
od events.
[Gallese et al. 1996]


Generazione di una rappresentazione interna
del movimento (simulazione), che pu avere
varie funzioni:

comprensione del signicato dellazione
osservata
leMura delle intenzioni dellindividuo che
agisce (Fogassi et al. 2005)



Fogassi et al. 2005
IPL (inferior parietal lobe)
neurons were studied when monkeys performed motor acts embedded in dierent
acNons and when they observed similar acts done by an experimenter.
Most motor IPL neurons coding a specic act (e.g., grasping) showed markedly dierent
acNvaNons when this act was part of dierent acNons (e.g., for eaNng or for placing).
Many motor IPL neurons also discharged during the observaNon of acts done by others.
Most responded dierenNally when the same observed act was embedded in a specic
acNon.

e.g.:
- intense ring when the monkey grasped a fruit to bring it to
its mouth.
- weaker response when the monkey grasped the food to place
it in a container.
- intense response when the monkey watched an
experimenter perform the grasp-to-eat gesture.
- weaker response to the grasp-to-place acNon.

These neurons not only code the observed motor

act but also allow the observer to understand the
agents intenNons.
 Neuroni specchio nelluomo
COME? Evidenze indireMe
EsperimenN di Npo neurosiologico
EEG - eleMroencefalograa: Registrazione del mu-rhythm (cfr. studi di Gastaut)
TMS - Transcranial magneNc sNmulaNon
brain-imaging
PET - Tomograa a emissione di positroni
fMRI- Risonanza magneNca funzionale

DOVE?
Osservazione di azioni compiute da altri comporta anche nelluomo la_vazione di un
network complesso di aree:
Aree visive (occipitali, temporali e parietali)
regioni corNcali eminentemente motorie
parte rostrale del lobulo parietale inferiore IPL
parte inferiore del giro precentrale
parte posteriore del giro frontale inferiore

Omologo di F5 nelluomo? AREA DI BROCA
 Area di Broca
Non unarea esclusivamente linguisNca!

Corrisponde alle aree di Brodmann 44 e 45.

- Area 44, Pars opercularis del giro frontale inferiore: in aggiunta alla rappresentazione del
linguaggio conNene, cos come F5 nella scimmia, le rappresentazioni motorie dei movimenN
della mano.
- Area 45, pars triangularis del giro frontale inferiore: a_vata da elemenN linguisNci, sia verbali
che segnaN.

Connessa allarea di Wernicke aMraverso il fascicolo arcuato
scimmia

uomo
Buccino et al. 2001
fRMI
AIM:
to localize brain areas that were acNve during the observaNon of acNons made
by another individual
to assess whether the observaNon of acNons made with dierent eectors
would acNvate specic parts of the premotor cortex in accord with the
somatotopic motor organizaNon of the region

STIMULI:
Object- and non-object-related acNons made with dierent eectors
(mouth, hand and foot) were presented.
mouth acNons:
biNng an apple
chewing
hand acNons:
reaching and grasping a ball or a liMle cup with the hand
mimicking these acNons without the object
foot acNons
kicking a ball
pushing a brake
mimicking these acNons without the object
FINDINGS:

ObservaNon of both object- and non-object-related acNons determined a
somatotopically organized acNvaNon of premotor cortex. The somatotopic
paMern was similar to that of the classical motor cortex homunculus.

During the observaNon of object-related acNons, an acNvaNon, also

somatotopically organized, was addiNonally found in the posterior parietal
lobe.

The mirror System is not restricted to the ventral premotor cortex, but
involves several somatotopically organized motor circuits.

LESPERIMENTO DIMOSTRA INOLTRE CHE:

Il sistema specchio nelluomo non circoscriMo alla mano.

Nelluomo anche losservazione di pantomine determina la_vazione

nei neuroni specchio. (cfr. Umilt et al. 2001)

Buccino et al. 2004

fRMI

AIM: to assess the corNcal areas acNve during the observaNon of

mouth acNons performed by humans and by individuals belonging
to other species (monkey and dog).

STIMULI: Two types of acNons were presented: biNng and oral

communicaNve acNons (speech reading, lip-smacking, barking).
FINDINGS:

The results showed that when the observed acNon is common to animals
and humans, there is a clear overlap between the acNvated areas, in spite
of the enormous dierences in the visual aspects of the observed sNmuli.
In contrast, during the observaNon of acNons that, like oral
communicaNve acNons, have a common goal, but are expressed
dierently in the three species, there is a clear dierence in the
distribuNon and extent of acNvaNons.

Taken together, the results of the present experiment suggest that

acNons made by other individuals may be recognized in dierent ways.
AcNons belonging to the motor repertoire of the observer are mapped on
his/her motor system. AcNons that do not belong to this repertoire appear
to be recognized essenNally based on their visual properNes.

AcNons that are not part of the motor repertoire of the observer and that
therefore cannot be reproduced appear to be recognized in nonmotor
terms. They are most likely understood based on visual descripNon of the
observed events and inferences of their consequences and/or goals.
 Neuroni specchio ed empaNa
Simulazione incarnata e rispecchiamento di emozioni e sensazioni.
Comprensione esperienziale delle emozioni di base, ad es. Disgusto
BASE BIOLOGICA DELLEMPATIA


Wicker et al. 2003

fRMI
STIMULI:
parNcipants inhaled odorants producing a
strong feeling of disgust.
the same parNcipants observed video clips
showing the emoNonal facial expression of
disgust.

Observing such faces and feeling disgust

acNvated the same sites in the anterior
insula and to a lesser extent in the
anterior cingulate cortex.

 Simulazione Incarnata
Il meccanismo funzionale che sta alla base del doppio paMern di a_vazione dei
neuroni specchio stato denito simulazione incarnata (embodied simula2on)

Nuova concezione delle a_vit mentali e della cognizione:

embodied (V. Gallese) / grounded (Barsalou)

da meccanismo di manipolazione di simboli astra_ e rappresentazioni, basato su

regole formali e processi a ci che emerge dallazione del corpo nel suo
contesto reale


Capacit perce_ve e motorie non sono separabili. Da esse si sono sviluppate
probabilmente anche le facolt superiori.

Il sistema sensomotorio di cui fano parte i neuroni specchio ha caraMere
mulNmodale, e non modulare!



 Riassumendo
Peculiarit dellarea F5: neuroni a_vi somatotopicamente in rapporto alla
funzione, ovvero sparano in concomitanza di a_ motori, non di singoli
movimenN.

2 popolazioni di neuroni nella scimmia nelluomo, con distribuzione

corNcale omologa:
Neuroni Canonici (Neuroni visuomotori che scaricano sia durante gli a_ motori,
sia alla presentazione di ogge_ 3D)

Neuroni Specchio (Neuroni visuomotori che scaricano sia durante gli a_ motori,
sia durante losservazione dello stesso aMo motorio compiuto da un altro agente)

SoMoclasse dei neuroni specchio audiovisivi

Funzione dei Neuroni specchio : comprensione dellazione + leMura dellintenzione
Peculiarit del sistema specchio nelluomo:
Non circoscriMo alla mano
Si a_va anche in caso di pantomina
Simulazione incarnata: le azioni che sono fuori dal repertorio della specie non hanno risonanza
motoria nellosservatore.
Neuroni specchio ed empaNa
Simulazione incarnata: quali implicazioni per la teoria della mente?
 Perch la scoperta dei neuroni specchio
dovrebbe interessare al linguista?

Filogenesi del linguaggio

Ontogenesi del linguaggio

Neuroni specchio, percezione e comprensione
linguisNca
Liv. Fonologico
Liv. Lessicale-semanNco
 MA PRIMA
Neuroni specchio nella scimmia: altri eeMori
Cfr. Buccino et al. 2001
il sistema specchio nelluomo non circoscriMo alla mano


Ferrari et al. 2003: NEURONI SPECCHIO DELLA BOCCA
Purpose of the experiments:
to study the responses of F5 neurons motorically coding mouth acNons to
the observaNon of mouth acNons made by another individual.

Results:
35% of mouth motor neurons discharged when the monkey observed
mouth acNons (mouth mirror neurons).
The majority of mouth mirror neurons (85%) discharged during the
observaNon of ingesNve mouth acNons, others (15%) during the
observaNon of communicaNve acNons.

INGESTIVE MOUTH-MIRROR NEURONS
The funcNonal properNes of ingesNve mouth-mirror neurons are very
similar to those of hand mirror neurons.
As hand mirror neurons, mouth mirror neurons require an interacNon
between an eector and an object in order to be triggered. The presentaNon
of an object alone or of a not goal-directed movement is insucient to
acNvate the neurons.
They show congruence between the eecNve observed and the eecNve
executed acNon.
these ndings extend the previous concept of the mirror system as a
system matching acNon observaNon and acNon execuNon from hand to
mouth acNons.

COMMUNICATIVE MIRROR NEURONS

communicaNve mirror neurons, unlike other mirror neurons, discharge in
response to acNons such as lip-smacking or tongue protrusion, acNons
that are not object-directed.
the eecNve motor acNon is not idenNcal to the eecNve observed one.
All studied communicaNve mirror neurons discharged during acNve
ingesNve acNons made by the monkey.

Why should ingesNon and communicaNon share a common neural substrate?

 Filogenesi del linguaggio

Come possibile che da tale sistema, cos streMamente legato
alloggeMo, si sia sviluppato un sistema aperto capace di
descrivere azioni e ogge_ senza necessariamente riferirvisi?

Rizzola_ & Arbib 1998
The observaNon/execuNon matching system provides a
necessary bridge from doing to communica2ng

The link between actor and observer becomes a link
between the sender and the receiver of each message.



 Filogenesi del linguaggio
The development of the human lateral speech circuit is a consequence of
the fact that the precursor of Brocas area was endowed, before speech
appearance, with a mechanism for recognizing acNons made by others.
This mechanism was the neural prerequisite for the development of
interindividual communicaNon and nally of speech.

HP: the mimeNc capacity inherent to F5 and Brocas area had the
potenNal to produce various types of closed systems related to the
dierent types of motor elds present in that area (hand, mouth and
larynx)

the rst open system to evolve en route to human speech was a manual
gestural system that exploited the observaNon and execuNon matching
system described earlier

gestural communicaNon vocalizaNons speech

 Filogenesi del linguaggio
Neuroni specchio della bocca [Ferrari et al. 2003]

HP: il sistema ingesNvo, controllato dalle aree motorie ventrali, si evoluto

nel sistema comunicaNvo

IngesNon and communicaNon share a common neural substrate
ethological and evoluNonary data that strongly suggest that these two
funcNons are strictly linked.
monkey communicaNve gestures (e.g. lip-smacking, lips protruded face)
are ritualizaNons of ingesNve acNons that are used for aliaNve purposes.
[] In other words, the knowledge common to the communicator and the
recipient of communicaNon about food and ingesNve acNon became the
common ground for social communicaNon. IngesNve acNons are the basis
on which communicaNon is built.

cfr. MacNeilage 1998: human vocal communicaNon derived from the cyclic
open-close mandibular alternaNon originally evolved for food ingesNon.
 Ontogenesi del linguaggio
(Capirci et al. 2005; Stefanini et al. 2009;
Congestr et al. 2010; PeMenaN et al. 2010)

Ruolo centrale del gesto nellontogenesi del
linguaggio, sia nelle primissime fasi dello sviluppo
comunicaNvo, sia nelle fasi successive.

ConNnuit tra le forme linguisNche e
prelinguisNche (ovvero tra azione-gesto-parola)

Progression from acNon to word through gesture

There is a conNnuity between the producNon of
the rst acNon schemes, the rst gestures and the
rst words produced by children. The relaNonship
between gesture and words changes over Nme.

1 anno: comunicazione intenzionale inizia mediante vocalizzazione e gesN dei_ci
(poinNng)

Gesto:
ProdoMo da solo
ProdoMo in combinazione con vocalizzazioni
ProdoMo in combinazione con 1word-uMerance

Cross modal combinaNon gesture/word combinaNon

Two-word stage
 Gestualit negli adulN
Contributo delle neuroscienze: approccio globale, integrato e
mulNmodale allanalisi dei processi comunicaNvi.

I gesN sono parte integrante del processo comunicaNvo

Il linguaggio un sistema integrato gesto-parlato
(cfr. Kendon 2004; McNeill 2000, 2005).
As a close examinaNon of the coordinaNon of gesture with speech
suggests, these two forms of expression are integrated, produced
together under the guidance of a single aim.
(Kendon, 2004: 2-3)

Nel linguaggio adulto i gesN sono coespressivi e sincronici rispeMo al

parlato: gesN e parole, pur usando modalit dierenN, sono inseparabili
(Unbreakable bond, McNeill 2005) ed esprimono ununica idea
soMostante, ovvero sono espressioni diverse dello stesso sistema.

Ruolo del gesto nellacquisizione di L1

 Gestualit negli adulN: case study
(Gagliardi, 2014)

Analisi della gestualit di sogge_ adulN ed anziani in un test di visual

confronta2on naming di azioni (daN della taratura della baMeria SMAAV,
Seman2c Memory Assessment on Ac2on verb)

Analisi linguisNca + analisi delle produzioni gestuali degli informanN

durante la somministrazione della baMeria.

Focus sui GesN rappresentaNvi (representa2onal) ovvero gesN che

forniscono una rappresentazione piMograca del signicato associato ad
un oggeMo o ad un evento. Denotano un preciso referente. Il contenuto
semanNco di base rimane relaNvamente stabile nelle diverse situazioni.

Ac2on gesture (o enactment): una parte del corpo compie un paMern di

azioni che hanno caraMerisNche in comune con il paMern di azioni che funge
da referente ;
size-shape gesture (o modeling / depic2on): il gesto richiama dimensione,
forma o caraMerisNche perce_ve dellevento o delloggeMo. (Ad esempio, le
mani assumono una congurazione che riproduce la forma delloggeMo a cui il
gesto si riferisce o eseguono un movimento per creare loggeMo nellaria.)
FINDINGS:

In accordo con quanto riportato in leMeratura, i signicaN convogliaN da
gesto e risposta vocale sono simili (match semanNco)

Lo stroke del gesto (ovvero la parte centrale del gesto, obbligatoria, dotata
di signicato, in cui concentrato lo sforzo energeNco) prodoMo
sincronicamente al parlato, con cui partecipa alla costruzione del senso
degli enunciaN.

La coespressivita di gesto e risposta vocale puo declinarsi in varie maniere:

talvolta il gesto sembra congurarsi come espressione parallela,
ridondante rispeMo al signicato espresso verbalmente, mentre in altre
occasioni precisa e restringe lapplicazione delleNcheMa lessicale.

Sebbene in alcuni casi il gesto sembri subentrare nel momento in cui vi

siano dicolt nel recupero lessicale il rapporto tra esso e il parlato non
mai di pura compensazione.

Probabile a_vazione somatotopica.

 Acquisizione ANpica: Sindrome dello
SpeMro AuNsNco
CARATTERISTICHE del DISTURBO:
incapacit di relazione interpersonale
mancanza di aMenzione
tendenza allisolamento
evitamento dello sguardo
considerazione dellaltro come un oggeMo

Gallese 2006
embodied mechanisms involving the acNvaNon of the sensory-motor system
seem to play a major role in social cogniNon.

all the decits can be explained as instanNaNons of intenNonal aMunement
decits produced by defecNve embodied simulaNon, likely underpinned by a
malfuncNoning of the mirror neuron systems.

Evidenze che vi sia un malfunzionamento nel sistema specchio:
EleMroencefalograa: gli individui auNsNci non mostrano soppressione del ritmo mu durante
losservazione delle azioni

DapreMo et al. 2006

fRMI

10 High-funcNoning children with auNsm and 10 typically developing children

AIM: to invesNgate neural acNvity during the imitaNon and observaNon of facial
emoNonal expressions

STIMULI: 80 faces expressing ve dierent emoNons:
anger
fear
happiness
neutrality
sadness

FINDINGS:
The evidence [] suggests that although both groups performed the imitaNon
task as requested, the neural strategies adopted by typically developing children
and those with ASD are quite dierent.
Reliable acNvity during imitaNon
of emoNonal expressions.

(a,b) AcNvity in bilateral pars
opercularis of the inferior frontal
gyrus is seen in the typically
developing group (a) but not in
the ASD group (b).

A between-group comparison (c)
 Typically developing children can rely upon a right hemispheremirroring
neural mechanisminterfacing with the limbic system via the insula
whereby the meaning of the imitated (or observed) emoNon is directly felt
and hence understood. In contrast, this mirroring mechanism is seemingly
not engaged in children with ASD, who must then adopt an alternaNve
strategy of increased visual and motor aMenNon whereby the internally
felt emoNonal signicance of the imitated facial expression is probably not
experienced.
ASD: Comprensione in termini cogniNvi, non esperienziali!

Examina2on of the rela2onship between

ac2vity in regions with mirror neuron
proper2es and symptom severity, as indexed
by childrens scores on the Au2sm Diagnos2c
Observa2on ScheduleGeneric (ADOS-G) and
the Au2sm Diagnos2c Observa2on Interview
Revised (ADI-R)

reliable negaNve correlaNons!!!
 Linguaggio nel bambino auNsNco
Brandi & Bigagli 2004
le caraMerisNche del disturbo portano a sospeMare la crucialit, anche per lo sviluppo
del linguaggio, di ci che cogniNvamente manca al bambino auNsNco: la capacit di
relazione intersogge_va, la capacit di rappresentare lo stato mentale altrui, la
capacit di imitare.

uno degli indicatori fondamentali della presenza del disturbo auNsNco lincapacit
nellesecuzione di giochi simbolici (tra 3 e 5 anni)

LINGUAGGIO (Brandi 2005)
ormai noto che lauNsmo, oltre ad essere propriamente un decit di Npo socio-
comportamentale, porN con s anche disturbi di caraMere streMamente linguisNco: a
seconda della gravit della sindrome possibile assistere ad un vero e proprio
muNsmo, oppure ad un notevole ritardo nello sviluppo del linguaggio (talvolta le
prime sillabe appaiono intorno agli oMo anni), o ancora, nel caso di uno sviluppo
linguisNco tardivo, esso pu manifestarsi aMraverso la produzione di stereoNpie


Disturbo complesso e di conseguenza eterogeneo per quanto riguarda la
manifestazione dei sintomi.
Varie tipologie di comportamento linguisNco. (vedi Brandi 2005)


STEREOTIPIE:
dissociazione tra forma e funzione nellacquisizione del linguaggio
dissociazione tra abilit sinta_co-semanNca e abilit pragmaNca

marcata asincronia tra lo sviluppo della funzione comunicaNva e
lacquisizione della grammaNca

Sono gli aspe_ pragmaNci del linguaggio a presentarsi sistemaNcamente
disturbaN nei sogge_ auNsNci

 Neuroni specchio e linguaggio:
livello fonologico
Fadiga et al. 2002
TMS

Subjects were instructed to listen carefully to a sequence of acousNcally presented

verbal and nonverbal sNmuli.
e.g.
WORDS: birra, carro, ferro, porro, terra
PSEUDO-WORDS: berro, furra, parro, vurro

Motor- evoked potenNals (MEPs) were recorded from opponens pollicis and
tongue muscles

FINDINGS:
during speech listening, there is an increase of motor-evoked potenNals recorded
from the listeners' tongue muscles when the presented words strongly involve,
when pronounced, tongue movements.

when an individual listens to verbal sNmuli there is an acNvaNon of the speech

related motor centres. Most interesNngly, they show also that this acNvaNon is
highly specic.
Watkins et al. 2003
TMS (Transcranial magneNc sNmulaNon) was applied to the face area of
primary motor cortex to elicit motor-evoked potenNals in the lip muscles.

AIM: to examine whether auditory percepNon of speech modulated the

excitability of the motor system underlying speech producNon.

EXPERIMENTAL CONDITIONS
(1) Speech condiNon: listening to speech (conNnuous prose) while viewing
visual noise.
(2) Non-verbal condiNon: listening to non-verbal sounds (e.g. glass breaking,
bells ringing, guns ring) while viewing visual noise.
(3) Lips condiNon: viewing speech-related lip movements while listening to
white noise.
(4) Eyes condiNon: viewing eye and brow movements while listening to white
noise.

FINDINGS: The results demonstrate that speech percepNon, either by

listening to speech or by visual observaNon of speech-related lip
movements, enhanced excitability of the motor units underlying speech
producNon.

 SinteNzzando:
Il sistema motorio si a_va durante percezione
del linguaggio (eeMori: lingua, labbra).

HP: Coinvolgimento del circuito dei neuroni

specchio nel processing linguisNco: lascoltatore
percepisce e decodica i suoni linguisNci
mappando linput sul repertorio di comandi
motori necessari per produrre qui suoni, che
anche lui possiede.

Simulazione incarnata della fonazione

 CoarNcolazione e il problema
dellinvarianza
COARTICOLAZIONE
A fundamental problem in speech percepNon is that the realizaNon of speech sounds
is highly context dependent. Successive speech sounds are produced by vocal tract
gestures that overlap temporally.
The consequence of coarNculaNon is that there is no one-to-one correspondence
between an acousNc event and the repertoire of phonemes or phoneNc features in
the language.

PROBLEMA DELLINVARIANZA: Invarianza del sistema che soggiace alla competenza
linguisNca vs. variabilit del parlato.
E comunque necessario un livello di rappresentazione fonologica?

Soluzione classica:
A view that is shared by many in the eld of speech percepNon is that speech is
perceived by exploiNng the same auditory mechanisms that analyse other classes of
environmental sounds. On the way to acNvaNng word forms in the mental lexicon
(lexical access), the acousNc informaNon gives way to an intermediate input
representaNon.
Phonemic level of representaNon
AlternaNve unit of percepNon (e.g. syllabe, stress unit)
 Motor theory of speech percepNon
ALTERNATIVA RADICALE!
Haskins Laboratories
Liberman et al. 1967; Liberman e Ma_ngly 1985, Galantucci et al. 2006

PrimiNvi linguisNci: i gesN arNcolatori.

Il linguaggio parlato viene percepito confrontando tali gesN arNcolatori

con il repertorio motorio dellascoltatore.

Percezione e produzione del linguaggio uNlizzano il medesimo repertorio

di primiNvi motori.

Carenza di evidenze sperimentali al momento della presentazione della

teoria. La scoperta dei neuroni specchio ha portato nuovi argomenN.

HP: A_vazione delle rappresentazioni motorie rilevanN fonologicamente.

Risonanza motoria durante la percezione.


Toni et al. 2008
the listener does not solve the invariance problem in the auditory
domain, but instead in the motor domain. The acousNc paMerns might be
dierent, but the arNculatory gestures that are needed to produce them
are the same.
A listener will understand speech by virtue of being a speaker.
The neuromotor commands provide the required invariance

Fadiga et al. 2002
A cogniNve translaNon into phonology is not necessary because the
arNculatory gestures are phonologic in nature. Furthermore, speech
percepNon and speech producNon processes use a common repertoire of
motor primiNves that, during speech producNon, are at the basis of
arNculatory gesture generaNon, while during speech percepNon, are
acNvated in the listener as the result of an acousNcally evoked motor
`resonance'.

SI PUO FARE A MENO DI UN LIVELLO DI RAPPRESENTAZIONE FONOLOGICO?


 Neuroni specchio e linguaggio:
livello semanNco
Buccino et al. 2005
TMS

AIM: to assess whether listening to acNon-related sentences modulates the

acNvity of the motor system.

By means of single-pulse TMS, either the hand or the foot/leg motor area in the
leq hemisphere was sNmulated in disNnct experimental sessions, while
parNcipants were listening to sentences expressing hand and foot acNons.
Listening to abstract content sentences served as a control. Motor evoked
potenNals (MEPs) were recorded from hand and foot muscles.

STIMULI: Fiqeen hand-acNon-related and 15 foot-acNon-related sentences were

presented. As control sNmuli, 15 sentences with verbs expressing an abstract
content were delivered. All sentences were in Italian. All acNon-related sentences
expressed a concrete acNon on an appropriate object (e.g., cuciva la gonna []).
All abstract content sentences expressed an abstract acNon on an appropriate
object (e.g., amava la patria []). All verbs were formed by three syllables and
were conjugated in the third person of the past tense.
Hand-acNon-related sentences:
Girava la chiave, Prendeva la tazza, scriveva il tema

Foot-acNon-related sentences:
Marciava sul posto, saltava il muro, pestava le foglie

Abstract content sentences:
Soriva il freddo, scordava la data, temeva la pena


Results showed that MEPs recorded from hand muscles were specically
modulated by listening to hand-acNon-related sentences, as were MEPs
recorded from foot muscles by listening to foot-acNon-related sentences.
Hauk et al. 2004

fMRI

AIM: The predicNon under invesNgaNon in the present study

concerns possible dierences between the corNcal acNvaNon
paMerns elicited by acNon words of dierent semanNc
subcategories and, more specically, their relaNon to motor areas

STIMULI: 50 words from each of the three semanNc subcategories

(i.e. acNon words referring to face, arm, leg acNons) were selected
and presented in a passive reading task to 14 right-handed
volunteers, while hemodynamic acNvity was monitored using
event-related fMRI.

To idenNfy the motor cortex in each volunteer individually,

localizer scans were also performed, during which subjects had to
move their leq or right foot, leq or right index nger, or tongue.
FINDINGS:
Our present results indicate that such acNon-related acNvaNon can
involve primary motor cortex and does not require a linguisNc task (e.g.,
naming) but is elicited by sNmulus words per se, even in a passive reading
task.

[] the paMern of corNcal acNvaNon elicited by an acNon word reects

the corNcal representaNon of the acNon to which the word refers.

Esperimento replicato molteplici volte! Ad esempio Te4aman5 et al. 2005

 In sintesi
Pulvermuller 2005
Hearing a word seems to be associated with acNvaNon of its arNculatory
motor program, and understanding an acNon word seems to lead to the
immediate and automaNc thought of the acNon to which it refers.

If acNon words are semanNcally related to the movements of the face or
arNculators, arm or hand, or leg or foot, the distributed neuronal ensembles
would include semanNc neurons in perisylvian (face-related words), lateral
(arm-related words) or dorsal (leg-related words) motor and premotor cortex.
Therefore, this semanNc somatotopy model of acNon words implies that
there are dierently distributed networks for the English words lick, pick
and kick
 Memoria SemanNca: denizione
La memoria semanNca, o conoscenza conceMuale, laspeMo della
memoria umana che corrisponde alla conoscenza generale del
signicato delle parole e di ogge_, fa_ e persone, senza
connessione ad un parNcolare tempo o luogo.

Seman2c memory is the memory necessary for the use of
language. It is a mental thesaurus, organized knowledge a person
possesses about words and other verbal symbols, their meanings
and referents, about rela2ons among them, and about rules,
formulas, and algorithms for the manipula2on of these symbols,
concepts and rela2ons. (Tulving 1972)

Vs. Memoria Episodica

Substrato neurale?
 Memoria SemanNca: correlaN neurali
Le posizioni teoriche correnN sulla memoria semanNca condividono
lidea che gran parte del contenuto della conoscenza conceMuale
sia collegata alla percezione ed allazione, e sia dunque un network
neurale distribuito, rappresentato somatotopicamente in regioni
cerebrali sovrapponibili o coincidenN con le aree senso-motorie

Due visioni contrapposte della rappresentazione neuroanatomica
della memoria semanNca (PaMerson et al. 2007):
distributed-only view: queste regioni cerebrali distribuite, e le
connessioni neurali che le collegano, esauriscono per intero la base
neurale della memoria semanNca
distributed-plus-hub view: le rappresentazioni provenienN dalle aree
senso-motorie e linguisNche sarebbero connesse e coordinate da un
hub amodale (o eteromodale) situato nel lobo temporale.
Distribuzione neuroanatomica dei network semanNci (PaMerson et al. 2007)

 Classi di parole nel cervello
Gli aMribuN semanNci connessi allazione e al movimento, generalmente
lessicalizzaN nei verbi, risiedono in struMure corNcali frontali, ovvero nelle
aree corNcali primariamente deputate alla programmazione ed esecuzione
motoria.
Il processing dei nomi avviene nellarea temporale anteriore e mesiale.
Per:
La generalizzazione non sembra estendersi al lessico nominale e verbale
astraMo (es. libert, e2ca).
La generalizzazione non sembra estendersi al linguaggio metaforico
(cfr. ad esempio la review di Aziz-Zadeh & Damasio 2008)
conce_ logici (es. IF, NOT)?

Lesioni focali e doppie dissociazioni
La scoperta di una doppia dissociazione tra nomi e verbi come forme lessicali
stata considerata a parNre dagli anni novanta unevidenza del faMo che la
classe grammaNcale sia un principio organizzaNvo della conoscenza lessicale
nel cervello (Hillis & Caramazza, 1995), rappresentata in network neurali
disNnN e separaN (Damasio & Tranel, 1993).

 Ipotesi alternaNve:
la dierenza di classe lessicale emerge a livello morfologico: non sono nomi e
verbi in s ad essere elaboraN in network separaN, ma i processi morfo-
sinta_ci che si applicano al lessico nominale e verbale sarebbero computaN in
reN neurali disNnte (Shapiro & Caramazza, 2003; Shapiro et al., 2006)
La dissociazione non tanto tra nomi e verbi come classi lessicali, quanto tra
parole riferite ad ogge_ e parole riferite ad azioni (Vigliocco et al. 2006, 2011)
 Fino a che punto possibile
generalizzare?
Toni et al. 2008
The discovery of mirror neurons in macaques and of a similar
system in humans has provided a new and ferNle neurobiological
ground for rooNng a variety of cogniNve faculNes.

Three claims that deal with the relaNonship between language and
the motor system:
Does language comprehension require the motor system?
Was there an evoluNonary switch from manual gestures to speech as
the primary mode of language?
Is human communicaNon explained by automaNc sensorimotor
resonances?

CRITICAL REVIEW



 We are not arguing against important links and
interacNons between language and motor systems in the
brain. For the semanNcs of certain word classes the acNon
system might be invoked. Clearly at the level of phoneNc
realizaNons the motor system plays a crucial role. This
could even be in language comprehension when under
condiNon of high predictability the predicted word forms
might actually be produced in the form of internal speech.
However, all this does not imply that the highly
complicated communicaNon system of natural language
can be fully reduced to sensorimotor properNes and the
contribuNon of sensorimotor areas.

Language goes beyond acNon. Without denying the

enormous importance of the discovery of mirror neurons,
their explanatory power for understanding human
communicaNon is limited.