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PCR - [Prodotti correlati](https://www.sigmaaldrich.com#prodotti) ## [](https://www.sigmaaldrich.com)Che cos'è il sequenziamento Sanger? Il sequenziamento Sanger, noto anche come "metodo di terminazione della catena", è un metodo per determinare la sequenza nucleotidica del DNA. Il metodo è stato sviluppato dal due volte premio Nobel Frederick Sanger e dai suoi colleghi nel 1977, da cui il nome di Sequenza Sanger. Per rivedere la struttura generale del DNA, si veda __Figura 2__. ## [](https://www.sigmaaldrich.com)Come funziona il sequenziamento Sanger? Il sequenziamento Sanger può essere eseguito manualmente o, più comunemente, in modo automatizzato tramite una macchina di sequenziamento (__Figura 1__). Ciascun metodo segue tre fasi fondamentali, come descritto di seguito. ![Sequenziamento Sanger automatizzato.](https://www.sigmaaldrich.com/deepweb/assets/sigmaaldrich/marketing/global/images/technical-documents/protocols/genomics/sequencing/sanger-sequencing_steps_process_diagram/sanger-sequencing_steps_process_diagram.png "Sequenziamento Sanger automatizzato.") __Figura 1.__Tre fasi fondamentali del sequenziamento Sanger automatizzato. ## [](https://www.sigmaaldrich.com)Fasi del sequenziamento Sanger Il sequenziamento Sanger prevede tre fasi principali. ### 1. Sequenza di DNA per la PCR a terminazione di catena La sequenza di DNA di interesse viene utilizzata come modello per un tipo speciale di [PCR](https://www.sigmaaldrich.com/IT/it/technical-documents/technical-article/genomics/pcr/routine-amplification) chiamata PCR a terminazione di catena. La PCR a terminazione di catena funziona come la PCR standard, ma con una differenza importante: l'aggiunta di nucleotidi modificati (dNTP) chiamati dideossiribonucleotidi (ddNTP). Nella fase di estensione della PCR standard, la DNA polimerasi aggiunge dNTP a un filamento di DNA in crescita catalizzando la formazione di un legame fosfodiestere tra il gruppo 3'-OH libero dell'ultimo nucleotide e il 5'-fosfato del successivo (__Figura 2__). Nella PCR a terminazione di catena, l'utente mescola un basso rapporto di ddNTP a terminazione di catena con i normali dNTP nella reazione di PCR. I ddNTP mancano del gruppo 3'-OH necessario per la formazione del legame fosfodiestere; pertanto, quando la DNA polimerasi incorpora un ddNTP a caso, l'estensione cessa. Il risultato della PCR a terminazione di catena è costituito da milioni o miliardi di copie oligonucleotidiche della sequenza di DNA di interesse, terminate a lunghezze casuali (n) da 5'-ddNTP. Nel sequenziamento Sanger manuale vengono allestite quattro reazioni di PCR, ciascuna con un solo tipo di ddNTP (ddATP, ddTTP, ddGTP e ddCTP). Nel sequenziamento Sanger automatizzato tutti i ddNTP sono mescolati in un'unica reazione e ognuno dei quattro dNTP ha un'unica etichetta fluorescente. ### 2. Separazione dimensionale mediante elettroforesi su gel Nella seconda fase, gli oligonucleotidi terminati a catena vengono separati per dimensione mediante elettroforesi su gel. Nell'elettroforesi su gel, i campioni di DNA vengono caricati in un'estremità di una matrice di gel e viene applicata una corrente elettrica; il DNA è carico negativamente, quindi gli oligonucleotidi saranno tirati verso l'elettrodo positivo sul lato opposto del gel. Poiché tutti i frammenti di DNA hanno la stessa carica per unità di massa, la velocità con cui gli oligonucleotidi si muovono sarà determinata solo dalle dimensioni. Quanto più piccolo è un frammento, tanto minore è l'attrito che subisce nel passaggio attraverso il gel e tanto più veloce sarà il suo movimento. Di conseguenza, gli oligonucleotidi saranno disposti dal più piccolo al più grande, leggendo il gel dal basso verso l'alto. Nel sequenziamento Sanger manuale gli oligonucleotidi provenienti da ciascuna delle quattro reazioni di PCR vengono fatti correre in quattro corsie separate di un gel. Questo permette all'utente di sapere quali oligonucleotidi corrispondono a ciascun ddNTP. Nel sequenziamento Sanger automatizzato tutti gli oligonucleotidi vengono eseguiti in un'unica elettroforesi su gel capillare all'interno della macchina di sequenziamento. ### 3. Analisi del gel e determinazione della sequenza del DNA L'ultima fase prevede semplicemente la lettura del gel per determinare la sequenza del DNA in ingresso. Poiché la DNA polimerasi sintetizza il DNA solo in direzione 5' - 3' partendo da un primer fornito, ogni ddNTP terminale corrisponderà a un nucleotide specifico nella sequenza originale (ad es, il frammento più corto deve terminare al primo nucleotide dall'estremità 5', il secondo frammento più corto deve terminare al secondo nucleotide dall'estremità 5', ecc.) Pertanto, leggendo le bande del gel dalla più piccola alla più grande, possiamo determinare la sequenza 5'-3' del filamento di DNA originale. Nel sequenziamento Sanger manuale l'utente legge tutte e quattro le corsie del gel in una sola volta, muovendosi dal basso verso l'alto, utilizzando la corsia per determinare l'identità del ddNTP terminale di ciascuna banda. Ad esempio, se la banda inferiore si trova nella colonna corrispondente a ddGTP, allora il frammento PCR più piccolo termina con ddGTP e il primo nucleotide dall'estremità 5' della sequenza originale ha una base di guanina (G). Nel sequenziamento Sanger automatizzato un computer legge ogni banda del gel capillare, in ordine, usando la fluorescenza per chiamare l'identità di ogni ddNTP terminale. In breve, un laser eccita i tag fluorescenti in ogni banda e un computer rileva la luce emessa. Poiché ciascuno dei quattro ddNTP è contrassegnato da un'etichetta fluorescente diversa, la luce emessa può essere direttamente collegata all'identità del ddNTP terminale. Il risultato è chiamato cromatogramma, che mostra il picco di fluorescenza di ciascun nucleotide lungo la lunghezza del DNA modello. ![Schema della struttura del DNA. Il DNA è una molecola composta da due filamenti che si avvolgono l'uno sull'altro a formare una doppia elica. Ogni filamento è costituito da una serie di molecole chiamate desossiribonucleotidi (dNTP).](https://www.sigmaaldrich.com/deepweb/assets/sigmaaldrich/marketing/global/images/technical-documents/protocols/genomics/sequencing/sanger-sequencing_dna-structure/sanger-sequencing_dna-structure.png "Schema della struttura del DNA. Il DNA è una molecola composta da due filamenti che si avvolgono l'uno sull'altro a formare una doppia elica. Ogni filamento è costituito da una serie di molecole chiamate desossiribonucleotidi (dNTP).") __Figura 2.__Schema della struttura del DNA. Il DNA è una molecola composta da due filamenti che si avvolgono l'uno sull'altro a formare una doppia elica. Ogni filamento è costituito da una serie di molecole chiamate desossiribonucleotidi (dNTP). Ogni dNTP contiene un gruppo fosfato, un gruppo zucchero e una delle quattro basi azotate \[adenina (A), timina (T), guanina (G) o citosina (C)]. I dNTP sono legati insieme in modo lineare da legami covalenti fosfodiestere tra lo zucchero di un dNTP e il gruppo fosfato del successivo; questo schema ripetuto zucchero-fosfato costituisce la spina dorsale zucchero-fosfato. Le basi azotate dei due filamenti separati sono legate insieme da legami idrogeno tra basi complementari per formare l'elica del DNA a doppio filamento. ## [](https://www.sigmaaldrich.com)Come leggere i risultati del sequenziamento Sanger La lettura corretta dei risultati del sequenziamento Sanger dipende da quale dei due filamenti di DNA complementari è di interesse e da quale primer è disponibile. Se i due filamenti di DNA sono A e B e il filamento A è di interesse, ma il primer è migliore per il filamento B, i frammenti in uscita saranno identici al filamento A. D'altra parte, se il filamento A è di interesse e il primer è migliore per il filamento A, l'uscita sarà identica al filamento B. Di conseguenza, l'output deve essere riconvertito in filamento A. Quindi, se la sequenza di interesse recita "TACG" e il primer è migliore per quel filamento, l'output sarà "ATGC" e, quindi, deve essere riconvertito in "TACG". Tuttavia, se il primer è migliore per il filamento complementare ("ATGC"), l'output sarà "TACG", che è la sequenza corretta. In breve, prima di iniziare, è necessario sapere a cosa si mira e come ci si arriva! Quindi, tenendo presente questo, ecco un esempio del primo esempio (TACG -> ATGC -> TACG). Se le etichette dei dideossinucleotidi sono T = giallo, A = rosa, C = blu scuro e G = azzurro, si otterranno le brevi sequenze primer-A, primer-AT, primer-ATG e primer-ATGC. Una volta separati i frammenti mediante elettroforesi, il laser leggerà i frammenti in ordine di lunghezza (rosa, giallo, blu chiaro e blu scuro) e produrrà un cromatogramma. Il computer convertirà le lettere, in modo che la sequenza finale sia il TACG corretto. ## [](https://www.sigmaaldrich.com)Sequenziamento Sanger vs. PCR Sanger Sequencing vs. PCR PCR Il sequenziamento Sanger e la PCR utilizzano materiali di partenza simili e possono essere usati insieme, ma nessuno dei due può sostituire l'altro. La PCR viene utilizzata per amplificare il DNA nella sua interezza. Sebbene si possano produrre accidentalmente frammenti di lunghezza variabile (ad esempio, la DNA polimerasi potrebbe cadere), l'obiettivo è duplicare l'intera sequenza di DNA. A tal fine, gli "ingredienti" sono il DNA bersaglio, i nucleotidi, il primer del DNA e la DNA polimerasi (in particolare la Taq polimerasi, che può sopravvivere alle alte temperature richieste dalla PCR). Invece, l'obiettivo del sequenziamento Sanger è quello di generare ogni possibile lunghezza di DNA fino all'intera lunghezza del DNA bersaglio. Per questo motivo, oltre ai materiali di partenza della PCR, sono necessari i dideossinucleotidi. Il sequenziamento Sanger e la PCR possono essere uniti quando si genera il materiale di partenza per un protocollo di sequenziamento Sanger. La PCR può essere utilizzata per creare molte copie del DNA da sequenziare. Avere più di un modello da cui lavorare rende il protocollo Sanger più efficiente. Se la sequenza target è lunga 1.000 nucleotidi e c'è una sola copia del modello, ci vorrà più tempo per generare i 1.000 frammenti marcati. Tuttavia, se ci sono diverse copie del templato, in teoria ci vorrà meno tempo per generare tutti i 1.000 frammenti marcati. ## Prodotti correlati Si è verificato un errore inatteso. 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