Classificazione dei recettori di membrana

I recettori di membrana, come abbiamo già detto nell’articolo precedente, trasducono il segnale portato da mediatori idrofilici che difficilmente passano la membrana cellulare. Sono proteine che si estendono attraverso tutto il doppio strato lipidico della membrana. Come i fosfolipidi, anche i recettori di membrana sono anfipatici, ossia possiedono regioni polari idrofile esposte sia sul versante extracellulare del plasmalemma sia su quello citoplasmatico, separate da segmenti idrofobici apolari che si inseriscono nel doppio strato fosfolipidico. La zona transmembrana di questi recettori è formata da Alfa-eliche composte da sequenze di 19-24 aminoacidi con gruppi laterali apolari. Generalmente, è la porzione idrofilica del recettore, posta nella zona extracellulare, ad interagire col neurotrasmettitore/farmaco, determinando una variazione conformazionale nella porzione del recettore posta nella zona citoplasmatica.

Struttura proteina di membrana. Fonte: Wikipedia.
Struttura proteina di membrana. Fonte: Wikipedia.

Recettori accoppiati a proteine G

Un tipo di recettore di membrana è quello rappresentato dai recettori accoppiati a proteine G. Quando attivati, essi producono secondi messaggeri. I secondi messaggeri rappresentano un sistema di amplificazione del messaggio del neurotrasmettitore. Sono recettori costituiti da una singola catena peptidica formata da sette domini transmembrana e con l’estremità N-terminale extracellulare e l’estremità C-terminale intracitoplasmatica. I siti di riconoscimento per i ligandi si trovano, a seconda del tipo di recettore, o sulla porzione N-terminale o nelle tasche formate dal raggruppamento delle porzioni transmembrana. La terza ansa, insieme ad un dominio della regione C-terminale, determina il sito di legame per la proteina G. Le proteine G (chiamate così perché legano il GTP) sono proteine formate da tre subunità; alfa, beta e gamma. Le subunità beta e gamma sono legate covalentemente tra loro. In condizioni di riposo, la subunità Alfa è legata ad una molecola di guanosina difosfato (GDP). Quando stimolato il recettore, la subunità Alfa rilascia il GDP e lega il GTP, mentre la subunità Beta-Gamma si stacca dal recettore. Si ritorna in una condizione di riposo quando la subunità Alfa, tramite la sua attività GTPasica, idrolizza la GTP in GDP e permette la sua riassociazione con la subunità Beta-Gamma riformando l’eterotrimero inattivo.

Recettori canale

Un altro tipo di recettore di membrana è il recettore canale (chiamato anche canale ionico controllato da ligando). Le subunità sono costituite da singole catene polipeptidiche che attraversano la membrana cellulare da due a sei volte e che formano un canale idrofilico in corrispondenza di regioni ricche di aminoacidi idrofobici (M1-M6). L’apertura del canale è causata dal legame del ligando ad un sito generalmente presente nella porzione extracellulare del recettore e produce un rapido e selettivo flusso ionico in entrata o in uscita. I recettori per i neurotrasmettitori acetilcolina, serotonina e acido glutammico sono permeabili al Sodio e al Calcio. Invece quelli per la glicina e il GABA sono permeabili al Cloro. I recettori canale possono essere suddivisi in diverse classi a seconda della natura e dalla tipologia di assemblaggio delle subunità.

  1. La prima classe è quella del recettore nicotinico dell’acetilcolina, del recettore GABAa del GABA, del 5-HT3 della serotonina e del GLY-R della glicina. Sono recettori eteropentameri cioè composti da un polipeptide che attraversa la membrana quattro volte. Ad esempio, il recettore nicotinico presenta due sub Alfa1, una sub Beta1, una sub Gamma1 e una sub Delta1.
  2. Alla seconda classe vi appartengono i recettori glutammatergici NMDA, AMPA e Kainato. Sono omotetrameri (o eterotetrameri) formati da subunità aventi quattro regioni transmembrana.
  3. La terza classe è composta dai recettori per i nucleotidi monofosfati ciclici (cAMP e cGMP). Le loro estremita N- e C- sono entrambe poste all’interno della cellula e son composti da subunità a sei regioni transmembrana.
  4. La quarta ed ultima classe è rappresentata dai recettori ionotropi dell’ATP, denominato P2X. Essi sono composti da 7 differenti subunità (P2X1-7) con due segmenti transmembrana.

Un tipico recettore canale è il recettore nicotinico per l’acetilcolina e per il GABA, entrambi formati da più subunità proteiche che attraversano la membrana plasmatica. Il recettore nicotinico per l’acetilcolina si trova nelle pieghe giunzionali della placca neuromuscolare nella membrana muscolare striata scheletrica, ai gangli del SNP (Sistema Nervoso Periferico). È un eteropentamero, a cinque subunità, in cui la subunità Alfa è presente in due copie mentre le altre sono presenti in copie singole. Il recettore classico per il GABA, quindi il GABAa, è un recettore ionotropo presente a livello postsinaptico ed è formato da due subunità Alfa, due subunità Beta ed una subunità Gamma. Come abbiamo già visto, al suo interno, presenta un canale per il cloro. Il GABA è agonista endogeno del recettore e si lega prevalentemente ad un sito posto sulla subunità Beta, mediando una modificazione allosterica che fa aprire il canale per il cloro.

Recettori per i fattori di crescita

Altra tipologia di recettori di membrana è quella dei recettori per i fattori di crescita. La maggior parte di questi recettori possiede un’attività enzimatica di tipo tirosino-chinasico e catalizza il trasferimento di gruppi fosfato dall’ATP ai gruppi idrossilici dei residui di tirosina presenti nelle proteine bersaglio. Un esempio classico di recettore per i fattori di crescita è quello per l’insulina*. Dal punto di vista funzionale, tali recettori sono organizzati in diversi domini:

  • Il dominio extracellulare comprende il sito di legame per il fattore di crescita. Esso presenta diverse combinazioni di motivi strutturali e, per questo motivo, è possibile raggruppare questo tipo di recettori in diverse sottofamiglie.
  • Il dominio transmembrana è costituito da circa 25 aminoacidi idrofobici che attraversano il doppio strato lipidico e funge da connessione tra il dominio extracellulare e quello intracellulare.
  • Il dominio iuxtumembrana comprende circa cinquanta aminoacidi e separa il dominio transmembrana da quello catalitico. Esso è sede di funzioni regolatorie ed è costituito da una regione ben conservata tra recettori appartenenti alla stessa sottofamiglia. Questa regione è coinvolta nella modulazione della trasduzione del segnale da parte di segnali extrarecettoriali.
  • Il dominio catalitico è responsabile dell’attività tirosino-chinasica. Esso è formato da 250 aminoacidi e ha un’elevata omologia fra tutti i recettori per i fattori di crescita.
  • La cosa C-terminale ha lunghezza e funzioni variabili e rappresenta il segmento che, nel recettore attivato, lega i trasduttori intracellulari del segnale.

Gran parte dei recettori per i fattori di crescita sono omodimeri (formati da due monomeri Alfa-Beta). Essi hanno un meccanismo secondo il quale il fattore si lega al dominio extracellulare.

Recettori per l’adesione cellulare

Esistono casi in cui il ligando ed il recettore sono due proteine transmembrana appartenenti a due cellule differenti. In questo caso, avviene una vera e propria comunicazione intercellulare che lega due o più cellule tra loro. I recettori che permettono questo meccanismo, infatti, vengono definiti “recettori per l’adesione cellulare”. L’interazione tra le due cellule viene detta omofilica se avviene tra due molecole di adesione dello stesso tipo. Invece, viene detta eterofilica se il legame avviene tra due molecole di adesione diverse o tra una molecola di adesione di una cellula e la matrice extracellulare di un’altra. Questi tipi di recettori possono essere raggruppati in grandi famiglie in base alla loro sequenza aminoacidica ed in base alle proprietà di legame. Le famiglie più caratteristiche sono le IgCAMs (gene per le immunoglobine), le integrine e le caderine.

  • I recettori della famiglia IgCAMs hanno regioni simili ai domini presenti nelle immunoglobine. Questi domini contengono serie di tratti idrofobici e idrofilici che formano una sequenza di foglietti Beta antiparalleli.
  • I recettori della famiglia delle integrine servono sia per l’adesione cellula-cellula sia per quella cellula-matrice extracellulare. Tutte le integrine mettono in contatto elementi del citoscheletro con proteine della matrice extracellulare.
  • I recettori della famiglia delle caderine sono formati da un dominio extracellulare N-terminale, un dominio transmembrana ed una regione citoplasmatica C-terminale. La regione extracellulare possiede cinque sequenze omologhe che, tramite il legame con ioni calcio, ne irrigidiscono la struttura. Il dominio intracellulare, invece, interagisce tramite proteine adattatrici con l’actina citoscheletrica.

Queste erano le grandi classi di recettori di membrana. A queste vanno aggiunte altre due classi (i recettori per le citochine e quelli con attività guanilato-ciclasica intrinseca) che sono di minor rilievo.

 

*L’insulina è un ormone proveniente dalle cellule beta del pancreas. La sua funzione è favorire l’utilizzazione del glucosio. Il recettore per l’insulina è costituito da due subunità Alfa e due subunità Beta. Ogni subunità Alfa si lega ad una subunità Beta formando due monomeri che, insieme, formano un dimero. La subunità Alfa presenta il sito di legame per l’insulina. Quando l’insulina si lega alla subunità Alfa, viene attivato il monomero alfa-beta che comporta l’autofosforilazione dei residui di tirosina presenti nell’altro monomero. Quando questo meccanismo è in corso, si l’attività tirosino-chinasica della parte più citosolica della catena beta.

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