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CRISPR-associated endonuclease Cas9 (EC 3.1.-.-) (St-Cas9)

 CAS9_STRTR              Reviewed;        1409 AA.
G3ECR1;
13-JUN-2012, integrated into UniProtKB/Swiss-Prot.
13-JUN-2012, sequence version 2.
10-OCT-2018, entry version 30.
RecName: Full=CRISPR-associated endonuclease Cas9 {ECO:0000255|HAMAP-Rule:MF_01480};
EC=3.1.-.- {ECO:0000255|HAMAP-Rule:MF_01480};
AltName: Full=St-Cas9;
Name=cas9 {ECO:0000255|HAMAP-Rule:MF_01480}; Synonyms=csn1;
Streptococcus thermophilus.
Bacteria; Firmicutes; Bacilli; Lactobacillales; Streptococcaceae;
Streptococcus.
NCBI_TaxID=1308;
[1]
NUCLEOTIDE SEQUENCE [GENOMIC DNA], FUNCTION IN PLASMID RESISTANCE,
EXPRESSION OF CRISPR3/CAS IN E.COLI, DISRUPTION PHENOTYPE, AND
MUTAGENESIS OF ASP-31; HIS-868; ASN-882 AND ASN-891.
STRAIN=DGCC7710;
PubMed=21813460; DOI=10.1093/nar/gkr606;
Sapranauskas R., Gasiunas G., Fremaux C., Barrangou R., Horvath P.,
Siksnys V.;
"The Streptococcus thermophilus CRISPR/Cas system provides immunity in
Escherichia coli.";
Nucleic Acids Res. 39:9275-9282(2011).
[2]
FUNCTION AS A DNA ENDONUCLEASE, COFACTOR, SUBUNIT, POSSIBLE
BIOTECHNOLOGY, DNA-BINDING, RNA-BINDING, EXPRESSION IN E.COLI, AND
MUTAGENESIS OF ASP-31 AND ASN-891.
STRAIN=DGCC7710;
PubMed=22949671; DOI=10.1073/pnas.1208507109;
Gasiunas G., Barrangou R., Horvath P., Siksnys V.;
"Cas9-crRNA ribonucleoprotein complex mediates specific DNA cleavage
for adaptive immunity in bacteria.";
Proc. Natl. Acad. Sci. U.S.A. 109:E2579-E2586(2012).
[3]
FUNCTION, ACTIVITY REGULATION, TRACRRNA-BINDING, RNA-BINDING, AND
EXPRESSION IN E.COLI.
STRAIN=DGCC7710;
PubMed=23535272; DOI=10.4161/rna.24203;
Karvelis T., Gasiunas G., Miksys A., Barrangou R., Horvath P.,
Siksnys V.;
"crRNA and tracrRNA guide Cas9-mediated DNA interference in
Streptococcus thermophilus.";
RNA Biol. 10:841-851(2013).
-!- FUNCTION: CRISPR (clustered regularly interspaced short
palindromic repeat) is an adaptive immune system that provides
protection against mobile genetic elements (viruses, transposable
elements and conjugative plasmids). CRISPR clusters contain
spacers, sequences complementary to antecedent mobile elements,
and target invading nucleic acids. CRISPR clusters are transcribed
and processed into CRISPR RNA (crRNA). In type II CRISPR systems
correct processing of pre-crRNA requires a trans-encoded small RNA
(tracrRNA), endogenous ribonuclease 3 (rnc) and Cas9. The tracrRNA
serves as a guide for ribonuclease 3-aided processing of pre-crRNA
(Probable). Cas9/crRNA/tracrRNA endonucleolytically cleaves linear
or circular dsDNA target complementary to the spacer yielding
blunt ends; Cas9 is inactive in the absence of the 2 guide RNAs
(gRNA). Cas9 recognizes a 3'-G-rich protospacer adjacent motif
(PAM, TGGTG in this organism) in the CRISPR repeat sequences to
help distinguish self versus nonself, as targets within the
bacterial CRISPR locus do not have PAMs. PAM recognition is also
required for catalytic activity. When the CRISPR3/cas system
consisting of cas9-cas1-cas2-csn2-CRISPR3 or just cas9-CRISPR3 is
expressed in E.coli it prevents plasmids homologous to spacers 1
or 2 from transforming. {ECO:0000269|PubMed:21813460,
ECO:0000269|PubMed:22949671, ECO:0000269|PubMed:23535272,
ECO:0000305}.
-!- COFACTOR:
Name=Mg(2+); Xref=ChEBI:CHEBI:18420;
Evidence={ECO:0000269|PubMed:22949671};
Note=Endonuclease activity on target DNA requires Mg(2+).
{ECO:0000269|PubMed:22949671};
-!- ACTIVITY REGULATION: Only has nuclease activity when bound to both
gRNAs (crRNA plus tracrRNA). {ECO:0000269|PubMed:23535272}.
-!- SUBUNIT: Monomer (Probable). Binds crRNA and tracrRNA.
{ECO:0000255|HAMAP-Rule:MF_01480, ECO:0000269|PubMed:22949671,
ECO:0000305}.
-!- DOMAIN: Has 2 endonuclease domains. The discontinuous RuvC-like
domain cleaves the target DNA noncomplementary to crRNA while the
HNH nuclease domain cleaves the target DNA complementary to crRNA.
{ECO:0000255|HAMAP-Rule:MF_01480}.
-!- DISRUPTION PHENOTYPE: Plasmid transformation is restored.
{ECO:0000269|PubMed:21813460}.
-!- BIOTECHNOLOGY: The simplicity of the Cas9-gRNAs RNA-directed DNA
endonuclease activity may be used to target and modify a DNA
sequence of interest.
-!- MISCELLANEOUS: This strain encodes 4 CRISPR-Cas systems; this is
CRISPR3.
-!- SIMILARITY: Belongs to the CRISPR-associated protein Cas9 family.
Subtype II-A subfamily. {ECO:0000305}.
-!- CAUTION: It is uncertain whether Met-1 or Met-22 is the initiator.
{ECO:0000305}.
-!- SEQUENCE CAUTION:
Sequence=AEM62887.1; Type=Erroneous initiation; Note=Translation N-terminally extended.; Evidence={ECO:0000305};
-----------------------------------------------------------------------
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EMBL; HQ712120; AEM62887.1; ALT_INIT; Genomic_DNA.
RefSeq; WP_024703962.1; NZ_QFLC01000001.1.
SMR; G3ECR1; -.
PRIDE; G3ECR1; -.
GO; GO:0003677; F:DNA binding; IEA:UniProtKB-KW.
GO; GO:0004519; F:endonuclease activity; IEA:UniProtKB-UniRule.
GO; GO:0046872; F:metal ion binding; IEA:UniProtKB-UniRule.
GO; GO:0003723; F:RNA binding; IEA:UniProtKB-KW.
GO; GO:0051607; P:defense response to virus; IEA:UniProtKB-UniRule.
GO; GO:0043571; P:maintenance of CRISPR repeat elements; IEA:UniProtKB-UniRule.
HAMAP; MF_01480; Cas9; 1.
InterPro; IPR028629; Cas9.
InterPro; IPR032239; Cas9-BH.
InterPro; IPR032237; Cas9_PI.
InterPro; IPR032240; Cas9_REC.
InterPro; IPR033114; HNH_CAS9.
InterPro; IPR003615; HNH_nuc.
Pfam; PF16593; Cas9-BH; 1.
Pfam; PF16595; Cas9_PI; 1.
Pfam; PF16592; Cas9_REC; 1.
Pfam; PF13395; HNH_4; 1.
TIGRFAMs; TIGR01865; cas_Csn1; 1.
PROSITE; PS51749; HNH_CAS9; 1.
1: Evidence at protein level;
Antiviral defense; DNA-binding; Endonuclease; Hydrolase; Magnesium;
Manganese; Metal-binding; Nuclease; RNA-binding.
CHAIN 1 1409 CRISPR-associated endonuclease Cas9.
/FTId=PRO_0000417879.
DOMAIN 792 949 HNH Cas9-type. {ECO:0000255|PROSITE-
ProRule:PRU01085}.
ACT_SITE 31 31 For RuvC-like nuclease domain.
{ECO:0000255|HAMAP-Rule:MF_01480}.
ACT_SITE 868 868 Proton acceptor for HNH nuclease domain.
{ECO:0000255|HAMAP-Rule:MF_01480}.
METAL 31 31 Magnesium 1. {ECO:0000255|HAMAP-
Rule:MF_01480}.
METAL 31 31 Magnesium 2. {ECO:0000255|HAMAP-
Rule:MF_01480}.
METAL 784 784 Magnesium 1. {ECO:0000255|HAMAP-
Rule:MF_01480}.
METAL 788 788 Magnesium 1. {ECO:0000255|HAMAP-
Rule:MF_01480}.
METAL 788 788 Magnesium 2. {ECO:0000255|HAMAP-
Rule:MF_01480}.
METAL 1011 1011 Magnesium 2; via pros nitrogen.
{ECO:0000255|HAMAP-Rule:MF_01480}.
MUTAGEN 31 31 D->A: No longer prevents plasmid
transformation. Target DNA
noncomplementary to the crRNA is not
cleaved. {ECO:0000269|PubMed:21813460,
ECO:0000269|PubMed:22949671}.
MUTAGEN 868 868 H->A: No longer prevents plasmid
transformation. Target DNA complementary
to the crRNA is not cleaved.
{ECO:0000269|PubMed:21813460}.
MUTAGEN 882 882 N->A: No longer prevents plasmid
transformation.
{ECO:0000269|PubMed:21813460}.
MUTAGEN 891 891 N->A: No longer prevents plasmid
transformation.
{ECO:0000269|PubMed:21813460,
ECO:0000269|PubMed:22949671}.
SEQUENCE 1409 AA; 163571 MW; 6A511E727C853AA6 CRC64;
MLFNKCIIIS INLDFSNKEK CMTKPYSIGL DIGTNSVGWA VITDNYKVPS KKMKVLGNTS
KKYIKKNLLG VLLFDSGITA EGRRLKRTAR RRYTRRRNRI LYLQEIFSTE MATLDDAFFQ
RLDDSFLVPD DKRDSKYPIF GNLVEEKVYH DEFPTIYHLR KYLADSTKKA DLRLVYLALA
HMIKYRGHFL IEGEFNSKNN DIQKNFQDFL DTYNAIFESD LSLENSKQLE EIVKDKISKL
EKKDRILKLF PGEKNSGIFS EFLKLIVGNQ ADFRKCFNLD EKASLHFSKE SYDEDLETLL
GYIGDDYSDV FLKAKKLYDA ILLSGFLTVT DNETEAPLSS AMIKRYNEHK EDLALLKEYI
RNISLKTYNE VFKDDTKNGY AGYIDGKTNQ EDFYVYLKNL LAEFEGADYF LEKIDREDFL
RKQRTFDNGS IPYQIHLQEM RAILDKQAKF YPFLAKNKER IEKILTFRIP YYVGPLARGN
SDFAWSIRKR NEKITPWNFE DVIDKESSAE AFINRMTSFD LYLPEEKVLP KHSLLYETFN
VYNELTKVRF IAESMRDYQF LDSKQKKDIV RLYFKDKRKV TDKDIIEYLH AIYGYDGIEL
KGIEKQFNSS LSTYHDLLNI INDKEFLDDS SNEAIIEEII HTLTIFEDRE MIKQRLSKFE
NIFDKSVLKK LSRRHYTGWG KLSAKLINGI RDEKSGNTIL DYLIDDGISN RNFMQLIHDD
ALSFKKKIQK AQIIGDEDKG NIKEVVKSLP GSPAIKKGIL QSIKIVDELV KVMGGRKPES
IVVEMARENQ YTNQGKSNSQ QRLKRLEKSL KELGSKILKE NIPAKLSKID NNALQNDRLY
LYYLQNGKDM YTGDDLDIDR LSNYDIDHII PQAFLKDNSI DNKVLVSSAS NRGKSDDFPS
LEVVKKRKTF WYQLLKSKLI SQRKFDNLTK AERGGLLPED KAGFIQRQLV ETRQITKHVA
RLLDEKFNNK KDENNRAVRT VKIITLKSTL VSQFRKDFEL YKVREINDFH HAHDAYLNAV
IASALLKKYP KLEPEFVYGD YPKYNSFRER KSATEKVYFY SNIMNIFKKS ISLADGRVIE
RPLIEVNEET GESVWNKESD LATVRRVLSY PQVNVVKKVE EQNHGLDRGK PKGLFNANLS
SKPKPNSNEN LVGAKEYLDP KKYGGYAGIS NSFAVLVKGT IEKGAKKKIT NVLEFQGISI
LDRINYRKDK LNFLLEKGYK DIELIIELPK YSLFELSDGS RRMLASILST NNKRGEIHKG
NQIFLSQKFV KLLYHAKRIS NTINENHRKY VENHKKEFEE LFYYILEFNE NYVGAKKNGK
LLNSAFQSWQ NHSIDELCSS FIGPTGSERK GLFELTSRGS AADFEFLGVK IPRYRDYTPS
SLLKDATLIH QSVTGLYETR IDLAKLGEG


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