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CME
Mirror neuron system
V. Rajmohan, E. Mohandas
Elite Mission Hospital, Thrissur, India
M i r r o r neur o n syst em i s a gr o up o f sp eci al i zed neur o ns
t hat mi r r o r s t he act i o ns and b ehavi o ur o f o t her s. The
i nvo l vement o f mi r r o r neur o n syst em (M NS) i s i mp l i cat ed
i n neur o co gni t i ve f unct i o ns (so ci al co gni t i o n, l anguage,
emp at hy, t heo r y o f mi nd ) and neur o p sychi at r i c d i so r d er s.
M NS d i sco ver y i s co nsi d er ed t o b e t he mo st i mp o r t ant
l and mar k i n neur o sci ence r esear ch d ur i ng t he l ast d ecad e.
MIRROR NEURON SYSTEM - DISCOVERY
The mi r r o r neur o ns w er e d i sco ver ed ser end i p i t o usl y b y
Gi aco mo Ri zzo l at t i and co l l eagues w hi l e w o r k i ng o n t he
gr asp r esp o nse o f macaq ues. They o b ser ved t hat a gr o up
o f neur o ns i n t he ar ea F5 o f t he p r emo t o r co r t ex t hat f i r es
w hen a macaq ue p er f o r ms an act i o n; al so d i schar ges w hen
i t o b ser ves t he same act i o n b ei ng p er f o r med b y ano t her
ani mal .
[ 1]
Sub seq uent r esear ch has el uci d at ed t he d i ver se
r egi o ns i nvo l ved i n t he M NS o f mo nk eys. Recent l y d i f f er ent
co r t i cal st r uct ur es have b een d escr i b ed as p ar t o f t he M NS
MNS i n m o n k eys
The ar ea F5 o f t he p r emo t o r co r t ex (p r emo t o r ar ea l o cat ed .
i n t he p o st er i o r b ank o f t he i nf er i o r ar cuat e sul cus and t he
co r t i cal co nvexi t y i mmed i at el y caud al t o i t ) i n mo nk eys has
t w o set s o f vi suo mo t o r neur o ns namel y t he cano ni cal and
t he mi r r o r neur o ns . The cano ni cal neur o ns (i n F5 b ank
r egi o n) r esp o nd t o p r esent at i o n o f an o b j ect w hi l e mi r r o r
neur o ns (i n F5 co nvexi t y) r esp o nd t o p er f o r mance o f an
act i o n and o b ser vat i o n o f an o b j ect d i r ect ed act i o n.
[ 2]
The
mi r r o r neur o ns ar e t r i gger ed b y any act i o n t hat i nvo l ves t he
t emp o r al sul cus (STS) and t he ar ea 7b (PF o f Vo n Eco no mo )
i n t he i nf er i o r p ar i et al l o b ul e (I PL). The STS co d es f o r a l ar ger
numb er o f mo vement s t han t he F5 neur o ns b ut l ack s mo t o r
p r o p er t i es (i .e., d o es no t d i schar ge w hi l e p er f o r mi ng t he
mo vement ). The ar ea 7b neur o ns i n I PL ar e het er o geneo us
and have a r o l e i n co d i ng senso r y st i mul i and r esp o nd t o
so mat o senso r y, vi sual o r b i mo d al st i mul i . I i n ad d i t i o n t o
t hi s, t hey al so have mo t o r p r o p er t i es and d i schar ge o n
act i o n o b ser vat i o n and p er f o r mance. The I PL r ecei ves i np ut s
f r o m t he STS and send s an i mp o r t ant o ut p ut t o t he vent r al
p r emo t o r co r t ex i ncl ud i ng ar ea F5 [ Fi gur e 1] .
[ 3,4]
MNS i n h um an s
Br ai n i magi ng st ud i es r eveal t hat act i o n o b ser vat i o n i n
humans act i vat es t he i nf er i o r f r o nt al gyr us (I FG), l o w er p ar t
o f t he p r ecent r al gyr us, t he r o st r al p ar t o f t he I PL and al so
t he t emp o r al , o cci p i t al and p ar i et al vi sual ar eas.
[ 3]
The f r o nt al
and t he p ar i et al mi r r o r neur o n r egi o ns ar e so mat o t o p i cal l y
o r gani zed . The act i vat i o n o f p ar s o p er cul ar i s o f t he I FG
r ef l ect s t he o b ser vat i o n o f d i st al hand and mo ut h act i o ns,
w her eas t he act i vat i o n o f t he p r emo t o r co r t ex r ef l ect s
p r o xi mal ar m and neck mo vement s [ Fi gur e 2] . The mi r r o r
neur o ns i n humans, unl i k e t ho se i n mo nk eys f i r e even
w hi l e o b ser vi ng meani ngl ess (i nt r ansi t i ve) mo vement s. The
SensoryInput
STS
IPL
i n humans.
i nt er act i o n b et w een a b i o l o gi cal ef f ect o r (mo ut h, hand et c.)
and an o b j ect . They ar e st i mul at ed b y t he o b ser vat i o n o f t he
exact same act i o n i nvo l vi ng t he ef f ect o r and o b j ect ( st r i ct l y
co ngr uent neur o ns) and al so b y act i o ns t hat ar e si mi l ar
b ut no t havi ng exact ef f ect o r -o b j ect i nt er act i o n ( b r o ad l y
co ngr uent neur o ns).
[ 3]
Ot her ar eas t hat f o r m p ar t o f t he M NS ar e t he sup er i o r
F5
F1
Correspondence: Dr. V. Raj mohan,
El i t e M i ssi on Hospi t al , Thri ssur, Indi a.
Motoroutput
E-mai l : emohandas53@gmai l .com
How to cite this article: Rajmohan V, Mohandas E. Mirror
neuron system. Indian J Psychiatry 2007;49:66-9.
Figure1:Basicmirrorneuroncircuitryinmonkey.F1:Primary
motorcortex,F5:Premotorcortex,IPL:inferiorparietallobule,
STS:superiortemporalsulcus.
Indian J Psychiatry 49(1), J an-Mar 2007 66
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Rajmohan et al.: Mirror neuron system
Mirror neuron system
Inferiorparictallobule
Primary
motor
cortex
Inferior
frontal
Acco r d i ng t o t hi s mo d el , act i o n und er st and i ng i nvo l ves
f i nd i ng a p r et end go al t hat w o ul d gener at e an act i o n
p l an i n t he o b ser ver s o w n mo t o r p l anni ng syst em. Thi s
i s t hen mat ched w i t h t he o b ser ved act i o n. I t i s p r esumed
t hat w hen t he si mul at ed mo t o r act i o n d o es no t mat ch t he
o b ser ved o ne, a new hyp o t hesi s i s gener at ed and t est ed
f o r co ngr uence w i t h t he o b ser ved act i o n. So act i o ns ar e
und er st o o d no t j ust i n t er ms o f t hei r o ut co mes, b ut al so
i n t er ms o f t he ment al st at es and esp eci al l y t he go al s, t hat
have gener at ed t hem.
[ 8]
I m i t at i o n
Basi c ci r cui t und er l yi ng i mi t at i o n co i nci d es w i t h t hat w hi ch
i s act i ve d ur i ng act i o n o b ser vat i o n. I mi t at i o n r eq ui r es a
p er f ect mat chi ng o f t he p er f o r med act i o n o nt o t he o b ser ved
Superiottemporalsulcus
Occipital
o ne. M i r r o r neur o ns ar e ab l e t o r eco gni ze t he act i o ns o f
lobe
o t her s and t he i nt ent i o n asso ci at ed w i t h t hem. So t hey can
co d e f o r l i k el y f ut ur e act i o ns o f o t her s, t her eb y o b ser ver s
Figure2:Mirrorneuronregionsinhumans ar e ab l e t o ant i ci p at e t he act i o ns o f o t her s.
[ 1,9]
o b ser vat i o n o f t r ansi t i ve act i o ns causes t he f i r i ng o f t he
Human i mi t at i o n i nvo l ves f l o w o f i nf o r mat i o n t hr o ugh
f r o nt al and t he t emp o r al no d es o f t he M NS w hi l e t hat o f
t he STS, I PL and t he I FG. STS p r o vi d es hi gher o r d er vi sual
i nt r ansi t i ve act i o ns r esul t i n t he f i r i ng o f t he f r o nt al no d e
p r o cessi ng o f o b ser ved act i o n w hi l e t he f r o nt o - p ar i et al M NS
co d es f o r t he go al o f t he act i o n and t he mo t o r p l an o n ho w
t o achi eve i t . The f r o nt o - p ar i et al syst em t hen send s co p i es
FUNCTIONS OF MNS
o f t hi s mo t o r p l an t o t he STS, w hi ch mat ches t he p r ed i ct ed
senso r y co nseq uence o f t he p l anned mo t o r act i o n w i t h
Act i o n un d er st an d i n g
t he vi sual d escr i p t i o n o f t he o b ser ved act i o n. The ef f er ent
Act i o n und er st and i ng i s t he f und ament al f unct i o n o f
co p i es t o t he STS o r i gi nat e f r o m t he vent r al sect o r o f t he
t he M NS. Each t i me t he ani mal o b ser ves a cer t ai n act i o n
p ar s o p er cul ar i s o f t he I FG, t he act i vi t y o f w hi ch i s sp eci f i c
b ei ng p er f o r med b y ano t her ani mal , t he mi r r o r neur o ns
t o i mi t at i o n.
[ 9]
r ep r esent i ng t he p er f o r mance o f t hat act i o n ar e act i vat ed .
The mi r r o r neur o ns t r ansf o r m vi sual o b ser vat i o n i nt o
Ther e i s a d i f f er ence d ur i ng i mi t at i ve l ear ni ng o f t ask s
k no w l ed ge.
[ 3]
St ud i es o n humans d ur i ng act i o n o b ser vat i o n
b et w een no vel t ask s and t he t ask s t hat ar e p ar t o f t he
have sho w n act i vat i o n o f t he I FG, t he I PL and a r egi o n w i t hi n
o b ser ver s mo t o r r ep er t o i r e. The f r o nt al and p ar i et al ar eas
t he STS. The p r ecent r al mo t o r co r t ex t ho ugh no t act i vat ed
ar e act i vat ed d ur i ng o b ser vat i o n o f a t ask t hat i s p ar t o f
b y act i o n o b ser vat i o n i s i nvo l ved i nd i r ect l y i n act i o n
o ne s r ep er t o i r e. The i mi t at i o n o f no vel act i o n i n ad d i t i o n
und er st and i ng as t hey have a r o l e i n mo t o r i mager y.
[ 7]
t o t he ab o ve r egi o ns i nvo l ves t he d o r so l at er al p r ef r o nt al
co r t ex (DLPFC) and t he co r t i cal ar eas r el evant t o mo t o r
The mai n hyp o t heses t o exp l ai n t he p heno meno n o f
p r ep ar at i o n namel y- d o r sal p r emo t o r co r t ex (PM d ), mesi al
act i o n und er st and i ng ar e t he vi sual hyp o t hesi s, d i r ect -
p r ef r o nt al co r t ex and t he sup er i o r p ar i et al l o b ul e. So t he
mat ch hyp o t hesi s and t he gener at e and t est mo d el .
b asi c i mi t at i o n l ear ni ng ci r cui t co nsi st s o f I FG, I PL, STS,
gyrus
o nl y.
[ 5,6]
Vi sual hyp o t hesi s i s b ased o n t he vi sual anal ysi s o f t he
ef f ect o r , t he o b j ect and o n t he co nt ext i n w hi ch t he act i o n
i s go i ng o n t o d r aw co ncl usi o ns as t o t he meani ng o f t he
act i o n. The neur al sub st r at es ar e vi sual ext r ast r i at e ar eas,
i nf er o t emp o r al l o b e and STS r egi o n. The d i r ect mat ch
hyp o t hesi s i s b ased o n t he map p i ng o f o b ser ved act i o n o n
hi s/her o w n mo t o r r ep r esent at i o n o f t he o b ser ved act i o n.
Ther ef o r e i t i nvo l ves a p r o cess o f o b ser vat i o n i nd uced mo t o r
r ep r esent at i o n, f o l l o w ed b y mat chi ng o f t hi s t o t he mo t o r
r ep r esent at i o n gener at ed d ur i ng act i ve o ut co me d i r ect ed
act i o n p er f o r med b y t he i nd i vi d ual (si mul at i o n). I f b o t h
t hese mo t o r r ep r esent at i o ns co r r esp o nd , i t l ead s t o act i o n
und er st and i ng.
[ 4]
A mo r e co mp l ex hyp o t hesi s suggest ed
f o r act i o n und er st and i ng i s t he gener at e and t est mo d el .
mo t o r p r ep ar at i o n ar eas and DLPFC [ Fi gur e 3] .
[ 9]
Sp eech an d l an guage
The p r esence o f mi r r o r neur o ns i n Br o ca s ar ea o f humans
suggest s t hat human l anguage may have evo l ved f r o m a
gest ur e p er f o r mance/und er st and i ng syst em. The t ask s l i k e
sp o nt aneo us sp eech and r ead i ng act i vat e t he hand mo t o r
ar ea and t he I FG, o n t he l ef t si d e.
[ 10,11]
So l anguage mi r r o r
neur o ns seem t o b e l at er al i zed t o t he l ef t si d e i nvo l vi ng
t he d o mi nant hand mo t o r co r t ex and t he hi gher l evel s o f
l anguage net w o r k .
[ 12]
Th eo r y o f m i n d
Theo r y o f mi nd (To M ) o r ment al i sat i o n i s t he ab i l i t y t o
Indian J Psychiatry 49(1), J an-Mar 2007 67
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Rajmohan et al.: Mirror neuron system
Table 1: Neural correlates of mirror neuron system,
theory of mind and empathy
DLPFC/PMd/MesialPFC/SPL
Mirror system Theory of mind Empathy
Imitative
Learning IFG IFG Premotor (IFG)
STS STS (Rt) STS (Rt)
IPL (Rt) IPL (Rt) IPL (Rt)
Occipital Occipital ACC
Primary motor cortex Med. PFC (inc ACC) VMPFC (inc OFC)
OFC Precuneus
Precuneus Somatosensory cortex
Amygdala
IFG - Inferior frontal gyrus, STS - Superior temporal sulcus, IPL- Inferior
parietal lobule, PFC - Prefrontal cortex, OFC - Orbitofrontal cortex, ACC
IFG/IPL/STS
Somatosensory cortex Amygdala
Imitation
Insula/LimbicSystem
Social
Insula
Posterior cingulate
Figure3:Mirrorneuronstructuresinimitation - Anterior cingulate cortex
r eco gni ze t hat so meo ne el se has a mi nd d i f f er ent f r o m o ne s d ysf unct i o nal M NS i n ASD. Neur o i magi ng st ud i es have
o w n. I t i nvo l ves t he ab i l i t y t o i nf er so meo ne el se s mi nd b y d emo nst r at ed l esser act i vat i o n o f Br o ad man s ar ea (BA)
f aci al exp r essi o n, t o ne o f vo i ce and no n-ver b al co mmuni cat i o n. 44/45, t he sup er i o r t emp o r al gyr us (BA 22), t he r i ght i nsul a
I t i nvo l ves t he ar ea co ncer ned w i t h act i o n i mi t at i o n, f ace and t he l ef t amygd al a.
[ 16]
A r ecent st ud y al so d emo nst r at ed
i mi t at i o n and i nt ent i o n und er st and i ng. The neur al st r uct ur es t he l ack o f M NS act i vi t y d ur i ng o b ser vat i o n and emo t i o nal
i nvo l ved i n To M i ncl ud e I FG, r i ght STS, r i ght I PL, med i al exp r essi o n i n chi l d r en w i t h ASD.
[ 17]
p r ef r o nt al co r t ex i ncl ud i ng t he ant er i o r ci ngul at e co r t ex
(ACC) o r b i t o f r o nt al co r t ex (OFC), p r ecuneus, so mat o senso r y The mi r r o r neur o ns may al so b e i nvo l ved i n t he d evel o p ment
co r t ex, amygd al a and t he o cci p i t al co r t ex. Ther ef o r e t he M NS o f d i so r d er s w i t h hyp er so ci al i t y l i k e Wi l l i am s synd r o me
i s co nsi d er ed i nt egr al t o t he t heo r y o f mi nd .
[ 13,14]
and Tur ner s synd r o me. The r el evance o f mi r r o r neur o ns
i n so ci al co gni t i o n may acco unt f o r d ysf unct i o nal M NS i n
So ci al co m m un i cat i o n an d em p at h y so ci al p ho b i a and aso ci al i t y o b ser ved i n schi zo p hr eni a.
[ 15]
So ci al co mmuni cat i o n and i d ent i f i cat i o n i nvo l ve i mi t at i o n.
The mo r e p eo p l e t end t o i mi t at e each o t her , t he mo r e The M NS al so p l ays an i mp o r t ant r o l e i n b o nd i ng and
t hey ar e ab l e t o d evel o p an emp at hi c r el at i o nshi p . So ci al at t achment . Dysf unct i o nal M NS may p l ay a r o l e i n ant i so ci al
mi r r o r i ng i nvo l ves t he i nt er act i o n o f t he co r e mi r r o r neur o n and b o r d er l i ne p er so nal i t y d i so r d er . The p at i ent s w i t h
syst em w i t h t he l i mb i c syst em. I magi ng st ud i es have sho w n b o r d er l i ne p er so nal i t y d i so r d er l ack t he ab i l i t y t o d i scer n
t hat o b ser vat i o n and i mi t at i o n o f f aci al emo t i o nal exp r essi o n t he ment al st at es o f sel f and o t her s. They have f r act ur ed
i nvo l ve t he f r o nt o -p ar i et al mi r r o r neur o n syst em, STS, i nsul a ear l y at t achment s l ead i ng t o a d ef i ci ency i n l ear ni ng o f
and t he l i mb i c syst em [ Fi gur e 3] .
[ 9]
t he co ncep t s l i k e secur e at t achment and t he cap aci t y t o
ment al i se.
[ 9,13]
Psycho t her ap eut i c p r o cesses i nvo l ve t he M NS
Emp at hy i s a p r o cess w hi ch i nvo l ves t he af f ect i ve shar i ng acco unt i ng f o r emp at hy and To M .
[ 13]
b et w een sel f and o t her s, ad o p t i ng t he p er sp ect i ve o f o t her s
and t he ab i l i t y f o r sel f agency and sel f r egul at i o n. The neur al CONCLUSION
co r r el at es emp at hy i ncl ud e I FG, r i ght STS, r i ght I PL, ACC,
vent r o med i al p r ef r o nt al co r t ex (VM PFC), so mat o senso r y The f asci nat i ng d i sco ver y o f M NS has gener at ed t r emend o us
co r t ex, amygd al a, p r ecuneus, i nsul a and t he p o st er i o r ent husi asm amo ng r esear cher s i n co gni t i ve neur o sci ence.
ci ngul at e. Thus emp at hy i nvo l ves a si gni f i cant i nt er act i o n
o f t he co r e M NS and i t s l i mb i c ext ensi o n
[ 9]
[ Tab l e 1] .
So ci al co gn i t i o n
So ci al co gni t i o n r ef er s t o t ho ught p r o cesses i nvo l ved i n
und er st and i ng and d eal i ng w i t h o t her s. I t i nvo l ves r egi o ns
t hat med i at e f ace p er cep t i o n, emo t i o nal p r o cessi ng, t heo r y
o f mi nd , emp at hy (esp eci al l y sel f r ef er ence) and w o r k i ng
memo r y. The M NS i nvo l vement i n med i at i ng emp at hy and
To M r ef l ect s i t s si gni f i cant r o l e i n so ci al co gni t i o n.
[ 15]
Mi r r o r n eur o n s an d n eur o p sych i at r i c d i so r d er s
Aut i sm and aut i sm sp ect r um d i so r d er s (ASD) ar e
char act er i zed b y so ci al i mp ai r ment , t he l ack o f To M and
d emo nst r ab l e d ef ect s i n i mi t at i o n sk i l l s. Thi s i mp l i es
The i nvo l vement o f M NS i n so ci al co gni t i o n, emp at hy
and To M has f uel l ed i nt er est i n exp l ai ni ng i t s r o l e i n ASD,
schi zo p hr eni a, p er so nal i t y d i so r d er s and p sycho t her ap eut i c
p r o cesses. The f ut ur e r esear ch may unr avel myst er i es
sur r o und i ng M NS.
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3. Rizzolatti G, Craighero L. The mirror-neuron system. Ann Rev Neurosci
2004;27:169-92.
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Indian J Psychiatry 49(1), J an-Mar 2007 68
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Source of Support: Nil, Conict of Interest:
Meister IG, Boroojerdi B, Foltys H, Sparing R, Huber W, Topper R. Motor
None declared
Announcement
TheLaunchofTheLancetSeriesonGlobalMentalHealthonSeptember3
rd
,2007
and
TheShortCourseonResearchMethodsinInternationalMentalHealth
(September4
th
to8
th
,2007)
ContactPersons:
ProfessorMartinPrince,InstituteofPsychiatry,KingsCollegeLondonandDr.VikramPatel,
LondonSchoolofHygieneandTropicalMedicine.
Pleasecontact:KCLsInternationalMentalHealthattheIoPcentre
atimh@iop.kcl.ac.ukorvisitwebpageat:www.iop.kcl.ac.uk/international
Indian J Psychiatry 49(1), J an-Mar 2007 69