Neurobiologia cunostintei:

Modelele neuroanatomice ale cunostintei:

1.

SRAA implicat in arousal este alcatuit din nucleii reticulari clasici din trunchiul cerebral, dar
includ si nucleii colinergici din trunchiul cerebral superior, nucleii noradrenergici, mai ales locus
coeruleus, proiecia histaminica din hipotalamusul posterior i probabil cai ascendente de dopamina si
serotonina din trunchiul cerebral. O mare parte din activitatea exercitat de aceste ci este mediata de
talamus, care poate fi considerat ca vrful SRAA, precum i un releu sinaptic critic pentru majoritatea
canalelor senzoriale i intracerebrale (Tononi i Koch, 2008 [220]; Tononi 2012 [222]). Rolul SRAA
implic, de asemenea, coordonarea cu talamusul, care prin colaterale axonale inhibitorii, inhiba
fasciculul reticular specific de informaii catre cortex. (Young i Pigott, 1999 [239]). Tomografia cu
emisie de pozitroni (PET) a demonstrat hipometabolism selectiv talamic i SRAA n timpul somnului
cu unde lente (Maquet, 2000 [137]) i anestezie (Alkire, Haier i Fallon, 2000 [3]).

2.

Amigdala neurobiologia a dovedit rolul ei in memoria de lunga durata (LTM), memoria de lucru
(WM) si atentie. Amigdala pare s medieze efectele componentei emotionale asupra LTM i atenie. De
asemenea, se pare c, faciliteaza performana cognitiv n timpul unei sarcini dificile i atunci cnd sunt
detectati stimuli motivaional relevanti (Arnsten i Li, 2005 [6]).Relaia dintre amigdala si sistemele
cognitive este bi-directionala, depinzand de link-uri funcionale corticolimbice. Astfel, noradrenalina i
dopamina pot juca un rol n relaia dintre amigdala i cunoaterea (cognition) superioar (Balleine i
Killcross, 2006 [16];. Kim et al, 2011 [107]). Apare evidenta nevoia de integrare intr-un model
cuprinztor, a bine-cunoscutei implicari a amigdalei n emoie, cu rolul su n cunoatere
superioara(Phelps, 2006 [166]). Amigdala asigura o funcie de vigilen generala, care are ca scop
pregtirea organismului pentru a face fa unor situaii dificile, solicitante, caz in care rspunsul vigilent
al amigdalei poate lua forma unor semnale neuromodulatorii fazice proiectate pe sistemul nervos
cognitiv, motor i / sau autonom, n timpul unei situaii ce solicita resurse metabolice suplimentare,
cum ar fi, o ameninare fizic sau o activitate mentala complexa (Schaefer i Gray, 2004 [185]). Pe de o
parte, amigdala este sensibila la stimuli care posed o valoare intrinsec de adaptare (de exemplu
stimuli emoionali, sau care genereaza teama etc). Pe de alt parte, amigdala este sensibila la stimuli a
cror valoare motivationala este determinat n mod arbitrar de scopul activitatii sau de anumite
solicitari (Richardson, ciudate i Dolan, 2004 [177]). Prin urmare, acest lucru sugereaza ca amigdala

Neurobiologia cunostintei:
trebuie s primeasc nu numai intrri senzoriale de baz, dar, de asemenea, intrri de la nivel nalt
legate de scopul sarcinii. Amigdala are un model remarcabil de bogat de conexiuni directe i indirecte
cu regiunile subcorticale i trunchiul cerebral, dar i cu regiunile neocorticale implicate n memoria de
lucru (WM) i controlul cognitiv, cum ar fi cortexul prefrontal (PFC) i cortexul cingular anterior
(ACC) (Maren, 2003 [138]; LaBar i Cabeza, 2006 [120]; Hirstein, 2011 [94]).

3.

Cerebelul cu rol dovedit in micare, mers, postura i echilibru. Cu toate acestea, unele studii au
sugerat o posibil implicare a cerebelului n cunoatere, prelucrarea emotiilor, cunotin i
comportament (Schmahmann, 2004 [194]; Schmahmann, Weilburg, i Sherman, 2007 [193]). Prin
aceste interconexiuni extinse: aferente (corticopontocerebelare) i eferente (cerebelotalamocorticale),
cerebelul poate primi informaii, i influena zone cerebrale corticale legate de cunoatere
(Schmahmann, 1997 [195]) [Figura 3]

Imagine reprodusa dupa http://www.benbest.com/

Au fost demonstrate interconexiuni semnificative ntre cerebel i subdiviziuni din cortexul prefrontal legate de
funciile executive (memorie de lucru, atenie, inhibarea comportamentului i luarea deciziilor), memorie verbala
si limbaj (Schmahmann i Pandya, 1995 [192]). De asemenea, exista teorii ce sustin implicarea proieciilor
aferente din parietal, cortexul temporal i occipital i sistemul limbic in integrarea de informaii senzoriale,
organizarea visuospatial, memoria vizuala, cunotin i controlul comportamentului i motivaie(Schmahmann
i Pandya, 1992 [191]) . Natura exact a implicrii cerebelului n procesele cognitive, este pn n prezent, mai
puin neleas. Anomalii ale structurii i funciei cerebelului au fost identificate neuroimagistic n anumite
tulburri psihiatrice (Konarski et al., 2006 [114]). Mai mult dect att, intervenii terapeutice farmacologice i
psihosociale pentru pacienii cu aceste tulburri au fost asociate cu modificri ale funciei cerebelului, sugernd
un rol important al acestuia, n diferite procese mentale, care sunt perturbate n anumite tulburri psihice (Strata,
2011[208]).
4.

Talamus - neuronii din talamusul central (nucleii intralaminari anterior i posterior, poriunile
paralaminare conexe ale nucleilor de asociere talamici - dorsal median, anterior ventral, ventral pulvinar
lateral i inferior) au specializri anatomice i fiziologice specifice care sprijin rolul lor esen ial n
funciile generale de atenie susinut, memorie de lucru i pregtire motorie. Aceste populaii neuronale
sunt interpuse ntre sistemul de excitare (arousal), care controleaz nivelul general de activitate talamic
i cortical, precum i sistemele de supraveghere frontale, care controleaza shift-ul premotor de
atenie i ajustarea nivelului de vigilen i alert (Parsuraman, 1999 [160]). n paralel, populatii

Neurobiologia cunostintei:
neuronale individuale din cadrul acestor agregate celulare sunt tintite de o varietate de relee senzoriale
dedicate care au evoluat, pentru a capta rapid atenia i a redireciona comportamentul (Krout, Belzer i
Loewy, 2002 [117]). Impreuna, aceste specializri anatomice sugereaz c muli neuroni din talamusul
central au rol in comportamentele directionate catre scop i in cunotin. Talamusul central primete
inervaie larga in cadrul sistemului de excitare (Praff, 2006 [164]) cu aferente neuronale colinergice
(Kolmac i Mitrofanis, 1999 [113]) cat i noradrenergica din locus ceruleus i aferente serotoninergice
de la rafeul medial (McCormick i Bal, 1997 [143]). Leziuni directe ale talamusului central pot produce
tulburri ale cunotinei la nivel global (Parvizi i Damasio, 2003 [161]). Dac leziunea este unilaterala,
se pot produce afectari hemispaiale sau stari modificate de cunostinta similare cu mania acuta sau
delirul. Leziuni bilaterale restrnse ale talamusului central pot produce coma acuta, care indic rolul
important pe care aceste structuri il joac in alerta, sau starea de veghe. Cu toate acestea, spre deosebire
de leziunile trunchiului cerebral care produc com n urma leziunilor focale, recuperarea cunotinei in
leziunile bilaterale centrale talamice n ceea ce privete comportamentul directionat spre scop i
abilitile de comunicare este foarte lent i incert (Schiff, 2008 [188]).

5.

Circuitele frontoparietale:

Daca cunotina fenomenal (phenomenal consciousness) se refer la aspectul subiectiv al experienei,

cunostinta de acces( access consciousness) se refer la aspectul obiectiv prin controlul direct al experienei
prin raionament, raportare, si aciune. Distincia ntre ele (Block, 1997 [23]), ar fi relevanta pentru a nelege
presupusul rol al zonelor frontoparietale n percepia contient. Phenomenal consciousness ar putea fi asociata
n principal cu activarea regiunilor senzoriale, iar access consciousness ar putea avea nevoie de implicare din
partea zonelor frontoparietale. Modelul specific de activare frontoparietala in timpul percepiei contiente poate
fi difereniat mai in detaliu n funcie de fazele i / sau tipul de prelucrare, de exemplu, tranziii perceptive fa de
percepie susinut. De asemenea exista teorii care sustin ca suprapunerea activarilor frontoparietale reflecta
procese cognitive comune, de tip atentional sau memorie de lucru.
6.

Cortexul prefrontal exista teorii contradictorii in ce priveste rolul in cunostinta, totusi cele mai recente
sustin c PFC joac un rol mai important n monitorizarea contienta dect n acumularea automat sau
non-contient de dovezi. Sugestia este c traseul contient de nivel superior corespunde reelei parietoprefrontal i zonelor corticale superioare asociate (Hirstein, 2011 [94]).

7.

Precuneus - a fost sugerat c activitatea precuneus n timpul strii contiente de odihn sprijin
procesarea conceptual care opereaz pe stocurile interne de informaii (semnale endogene), mai
degrab dect funciile de "percepie" (surse de informaii externe de creier). Cu alte cuvinte, acest
domeniu pare s contribuie la auto- prelucrarea de ganduri pe care oamenii o experimenteaza n
timpul odihnei constiente (Cavanna, 2007 [33]). O interaciune ntre precuneus i cortexul prefrontal a
fost postulata n stri de cunostinta caracterizate de un nivel ridicat de reflectare a contiinei de sine.
(Kjaer et al., 2002[109]; Cavanna and Trimble, 2006[32]).

Modele si teorii neuronale:

1. Teoria dendritelor apicale - Activitatea sincrona n grupurile de dendrite apicale produce unde
electromagnetice (EM) care pot radia spre exterior, iar n cazul n care acestea sunt suficient de
puternice pentru a ajunge la suprafaa pielii capului pot fi msurate ca EEG. Cmpul electromagnetic a
fost propus ca substrat fizic al cunotinei (McFadden, 2000 [144]; Pockett, 2000 [167]). Curenii
fluctuanti care produc cmpuri EM sunt produsi de mai multe surse, printre care: activitatea membranei
axonilor, dendritelor i somei neuronilor din regiunile corticale si subcorticale. Conform acestei teorii
cunotina de moment este reprezentata de cmpul general format din toate aceste domenii EM
individuale.
2.

Activitatea N-metyl D Asparat-ului

Neurobiologia cunostintei:
Apariia strii de cunostinta depinde n mod critic de o anumit categorie de procese computerizate,
care sunt mediate de synapse NMDA. Argumentele n favoarea ipotezei se bazeaz pe ideea c
activitatea receptorului NMDA este capabil s formeze reprezentari n creier i toate anestezicele
generale inhiba activitatea receptorilor NMDA. Prin urmare, activitatea receptorului NMDA este
esenial pentru cunostinta (Gambrill i colab., 2011 [77]).
3.

Modelul ansamblurilor neuronale:

Un ansamblu de celule neuronale este definit ca o structur difuz cuprinznd celule din cortex i diencefal,
capabil s acioneze rapid ca un sistem nchis i sa faciliteze functionarea altor astfel de sisteme (Hebb, 1949
[92]). Au fost stabilite astfel de ansambluri, observand ca 10 de milioane de neuroni isi pot sincroniza activitatea
n mai putin de 230 ms si s-a postulat ca fiind eseniale pentru producerea de stri contiente. n teorie neuronala
(Libet, 1993 [130]), cunotina este asociata cu activitile neuronale care persista pentru o perioad suficient de
lung, cu o durat minim de 500 ms. n sinapsele de ansambluri neuronale, cunoscute sub numele de sinapse
Malsburg, puterea sinapselor crete cnd exist o corelaie puternic ntre activitatea pre- i postsinaptic (Crick,
1984 [49]). Aceste configuraii tridimensionale de ansambluri de mari dimensiuni din creier, care nu respecta
limitele anatomice convenionale, se vor corela cu diferite grade de cunostinta n fiecare moment (Greenfield,
1997 [86]). Un studiu recent sprijina acest model de asamblare neuronala (Koch, 2005 [112]; Overgaard i
Morgensen, 2011 [157], Jansen i Overgaard, 2011 [101]).
4.

Modelul dialogului talamo-cortical:

Teoria descrie centre pe circuitul thalamocortical care implic nucleii intralaminari, care con in celule care
conecteaza activitatea trunchiul cerebral (in strile generale de veghe i vigilen) la diferite zone ale
cortexului (Bogen, 1995 [24]), distribuite pe scar larg. Alii au propus ca structurile talamice i corticale
(mpreun cu relaia lor cu nucleii reticulati din trunchiul cerebral) sunt condiii necesare i suficiente pentru
existena unei stari de cunostinta de veghe, iar proprietile recurente de "re-intrare" ale circuitului
thalamocortical par a fi esentiale (Baars et al, 2003 [9];.. Seth et al, 2005 [197]).
5.

Teoria activitatii neuronale cerebrale bazale:

S-a observat ca i n absena unor sarcini cognitive sau de stimulare senzorial, activitatea cerebrala
bazala n zonele implicate n geneza percepiei contiente fluctueaz continuu n creierul uman (Aglioti
i Candidi, 2011 [1]). Mai mult dect att, datele de cercetare sugereaza ca auto-contientizare i
constientizarea externa nu sunt concurente, ci ar fi mai degrab n competiie n timpul procesului de
percepie senzorial. Luate n ansamblu, aceste date sugereaz c fluctuaiile in activitatea cerebrala
bazala sunt susceptibile de a forma coninutul de "flux al contiinei" noastre n curs de desfurare ntrun mod decisiv. Cu toate acestea, originea i fiziologia acestor fluctuaii rmne un mister, i va face
obiectul unor studii viitoare (Boly et al, 2008 [27];.. Boly et al, 2011 [26]).

Teoria proto-panpsihista a cunostintei:

Conform acesteia sistemele contiente sunt cele care conin dou reele care interacioneaz activ, unul
pentru construirea de cunotine, i cealalt pentru generarea de sentimente. Noutatea este modul n care
este conceput sistemul de sentimente: ca un sistem care funcioneaz cu modele wavelike, iar fiecare
tip de val corespunde unui sentiment. Centrul care iniiaz sentimentele se propune a fi reeaua
astrogliala (Pereira Jr. i Furlan, 2010 [246]; Pereira Jr, Pereira i Furlan, 2010 [247]), reflectnd stari
ale ntregului corp, n procesul de interaciune cu lumea. n acest sens, studii recente fac legatura intre
undele de calciu din reeaua de astrocite cu integrarea informaiilor in creier (Navarrete et al., 2012
[248]), starea de contien (Thrane et al., 2012 [249]) i bolile neurologice cu impact asupra
cunontinei, cum ar fi boala Alzheimer (Furman i colab., 2012 [250]).
Reeaua astrogliala primete semnale de la ntregul corp prin intermediul neuronilor, fluxului de snge
i lichidul cefalorahidian (de exemplu, n sistemul vestibular). Aceast stare a ntregului corp este
comparata dinamic cu informaiile cu privire la existenta organismului n mediul exterior, aa cum este
reprezentat de reeaua neuronal. Rezultatul acestei comparaii este un sentiment de moment care
raporteaza informaiilor primite la starea sistemului. Acest sentiment impacteaza intreg organismul, prin
semnale neuro-endocrin-imune - inclusiv activiti ale inimii si sistemului nervos. Prin urmare,
contiina este conceputa ca prelucrarea de informaii despre corp i lume, construirea de cunotine i

Neurobiologia cunostintei:
investirea lor cu sentimente care influeneaz procesarea i, prin urmare, aciunile individului care
trieste n lume.

Neuromodulatorii:
Sistemul colinergic este unul dintre cele mai importante sisteme de neurotransmitatori la nivelul
creierului i controleaza activitile care depind de atenia selectiv, component esenial a cunostintei.
Dovezi psihofarmacologice i patologice susin conceptul de "component colinergica" a cunostintei. Pe
baza constatarii ca activarea corticala se mentine in cadrul somnului REM, in absenta activarii
neuronale mono-aminergice (serotoninergice sau noradrenergice) dar in prezenta descarcarilor constante
din neuronii colinergici ai nucleului bazal, s-a ajuns la concluzia ca sistemul colinergic ascendent este
suficient pentru mentinerea neocortexului intr-un mod operativ. Se pare insa ca sunt necesare aciuni
combinate ale ACh i glutamatului n acest sens. Teoriile recente ale rolului ACh includ posibilitatea ca
aceasta afecteaz procesele discriminatorii; creste rata semnal-zgomot, moduleaz eficien a de
prelucrare corticala a informaiilor senzoriale i de asociere, controleaz receptia stimulilor i evaluarea
nivelul lor de semnificaie, modific reacia corticala n ceea ce privete relevana i noutatea, i
limiteaz coninutul fluxului contient. Droguri care antagonizeaz receptorii muscarinici induc
halucinaii i reduc nivelul de cunostinta, n timp ce receptorul nicotinic este implicat n mecanismul de
aciune al anestezicelor generale (prin inhalare). In bolile degenerative ale creierului, modificri ale
strii de contien sunt asociate cu deficite regionale n sistemul colinergic. Reteaua vasta colinergica
din trunchiul cerebral care include proiecii convergente la talamus, pare a fi amplasata strategic pentru
a genera i integra constienta. [251]
Printre diferitele pri ale neuronului , dendritele sunt probabil cei mai buni candidati pentru a fi cheia
pentru functia cognitiva superioara , deoarece pot integra un numr mare de intrri . Membrana
neuronala este site-ul iniial de rspuns pentru intrri de la alti neuroni , dar ceea ce se afl dincolo de
membrana neuronal controleaz nivelul sinaptic de rspuns prin compararea intrrilor noi cu
informaiile stocate n memorie . Dendritele sunt imbogatite cu microtubuli si proteine asociate
microtubulilor (MAP) , al caror rol nu se cunoaste inca pe deplin. Acumularea de dovezi sugereaza ca
microtubulii i MAP-urile joac un rol critic n memorie i contiin , precum i n transportul
neuronal. Proteina - 2 asociata microtubulilor ( MAP2 ) este o protein citoscheletala specifica
dendritelor , care , de asemenea, acioneaz ca o molecul de transducie a semnalului , mediind
rspunsurile chimice interne ca urmare a eliberrii sinaptice de neurotransmitatori glutamat i
acetilcolina . MAP2 i microtubulii se leaga mpreun pentru a forma o matrice care stocheaz
memoria. Informaia stocata n matricea de microtubuli este apoi accesata la eliberarea anumitor
neurotransmitori , cum ar fi acetilcolin i glutamat . Acetilcolina controleaz nivelul de con tien ,
n principal prin receptorul muscarinic determinand activarea consecutiva a kinazelor PKC i CaMKII ,
ambele fosforiland MAP2 cu rol in memorie . Prin activarea de catre neurotransmitatori , microtubulii
pot evalua intrrile sinaptice actuale n contextul activitii sinaptice anterioare , i apoi sa creasca
transportul de anumite molecule legate de nvare la sinapse . Pentru aparitia activitatii con tiente este
necesara co- activare a unui numr mare de sinapse si transportul n mas de proteine sinaptice eseniale
, i este posibil ca quantum computation sa joace un rol n dirijarea coerenta a acestui transport . O
teorie anterioara sustine ca acetilcolina faciliteaz calculele cuantice n microtubuli prin fosforilarea
MAP2. O terorie recenta propune ca modelul de legare a MAP2 de microtubuli formeaz un contur pe
baza de gel, care reprezint informaii stocate de ctre mecanismul de nvare i ofer o baz fizic
pentru aceste informatii. Cnd MAP2 este fosforilat, acest contur pe baza de gel se extinde propagand
informaia longitudinal pe microtubul, i tangenial, la microtubulii vecini. n aceste dou moduri,
microtubulii activati fizic asigura un rspuns coerent care acoper mai multe sinapse moduland intrri
noi n contextul informaiilor stocate n memorie.
[252]
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