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Q6ZNE5

- BAKOR_HUMAN

UniProt

Q6ZNE5 - BAKOR_HUMAN

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Protein

Beclin 1-associated autophagy-related key regulator

Gene

ATG14

Organism
Homo sapiens (Human)
Status
Reviewed - Annotation score: 5 out of 5- Experimental evidence at protein leveli

Functioni

Required for both basal and inducible autophagy. Determines the localization of the autophagy-specific PI3-kinase complex. Plays a role in autophagosome formation and MAP1LC3/LC3 conjugation to phosphatidylethanolamine. Promotes BECN1 translocation from the trans-Golgi network to autophagosomes. Enhances PIK3C3 activity in a BECN1-dependent manner.4 Publications

GO - Biological processi

  1. autophagic vacuole assembly Source: UniProtKB
  2. endosome to lysosome transport Source: MGI
  3. positive regulation of autophagy Source: InterPro
Complete GO annotation...

Keywords - Biological processi

Autophagy

Names & Taxonomyi

Protein namesi
Recommended name:
Beclin 1-associated autophagy-related key regulator
Short name:
Barkor
Alternative name(s):
Autophagy-related protein 14-like protein
Short name:
Atg14L
Gene namesi
Name:ATG14
Synonyms:KIAA0831
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
ProteomesiUP000005640: Chromosome 14

Organism-specific databases

HGNCiHGNC:19962. ATG14.

Subcellular locationi

Cytoplasm. Endoplasmic reticulum membrane; Peripheral membrane protein. Preautophagosomal structure membrane; Peripheral membrane protein
Note: Cytosolic under nutrient-rich conditions. Following autophagy stimuli, such as starvation or rapamycin induction, predominantly detected in cytoplasmic foci, identified as isolation membranes and autophagosomes. Accumulates on highly curved PtdIns3P enriched autophagic membrane via its BATS domain to sense and maintain membrane curvature (By similarity). Localizes also to discrete punctae along the ciliary axoneme and to the base of the ciliary axoneme (By similarity).By similarity

GO - Cellular componenti

  1. autophagic vacuole Source: UniProtKB
  2. axoneme Source: UniProtKB
  3. cytoplasm Source: HPA
  4. endoplasmic reticulum Source: UniProtKB-KW
  5. nucleus Source: HPA
  6. phagocytic vesicle Source: Ensembl
  7. pre-autophagosomal structure membrane Source: UniProtKB
Complete GO annotation...

Keywords - Cellular componenti

Cytoplasm, Endoplasmic reticulum, Membrane

Pathology & Biotechi

Mutagenesis

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Mutagenesisi43 – 431C → A in Atg14L4C4A; fails to localize to the endoplasmic reticulum; when associated with A-46, A-55 and A-58. 1 Publication
Mutagenesisi46 – 461C → A in Atg14L4C4A; fails to localize to the endoplasmic reticulum; when associated with A-43, A-55 and A-58. 1 Publication
Mutagenesisi55 – 551C → A in Atg14L4C4A; fails to localize to the endoplasmic reticulum; when associated with A-43, A-46 and A-58. 1 Publication
Mutagenesisi58 – 581C → A in Atg14L4C4A; fails to localize to the endoplasmic reticulum; when associated with A-43, A-46 and A-55. 1 Publication

Organism-specific databases

PharmGKBiPA165478560.

PTM / Processingi

Molecule processing

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Chaini1 – 492492Beclin 1-associated autophagy-related key regulatorPRO_0000050774Add
BLAST

Amino acid modifications

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Modified residuei29 – 291Phosphoserine2 Publications
Modified residuei416 – 4161Phosphoserine1 Publication
Modified residuei429 – 4291Phosphothreonine1 Publication

Keywords - PTMi

Phosphoprotein

Proteomic databases

MaxQBiQ6ZNE5.
PaxDbiQ6ZNE5.
PRIDEiQ6ZNE5.

PTM databases

PhosphoSiteiQ6ZNE5.

Expressioni

Gene expression databases

BgeeiQ6ZNE5.
CleanExiHS_KIAA0831.
GenevestigatoriQ6ZNE5.

Organism-specific databases

HPAiCAB037242.
HPA053515.

Interactioni

Subunit structurei

Component of the autophagy-specific PI3-kinase complex I composed of ATG14, BECN1, PIK3C3 and PIK3R4, but not UVRAG, nor KIAA0226/Rubicon. UVRAG and ATG14/Barkor form mutually exclusive complexes with BECN1 through direct competition. The complex containing ATG14 up-regulates autophagy, while the one containing Rubicon down-regulates autophagy (By similarity). Interacts with PIK3CB (By similarity). Interacts with BECN1P1/BECN2.By similarity5 Publications

Binary interactionsi

WithEntry#Exp.IntActNotes
BECN1Q1445711EBI-2690371,EBI-949378

Protein-protein interaction databases

BioGridi116531. 13 interactions.
DIPiDIP-48651N.
IntActiQ6ZNE5. 7 interactions.
MINTiMINT-8304673.

Structurei

3D structure databases

ProteinModelPortaliQ6ZNE5.
ModBaseiSearch...
MobiDBiSearch...

Family & Domainsi

Region

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Regioni43 – 5816Cysteine repeatsAdd
BLAST
Regioni413 – 49280BATSBy similarityAdd
BLAST

Coiled coil

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Coiled coili71 – 180110Sequence AnalysisAdd
BLAST

Domaini

The coiled-coil domain is required for BECN1- and PIK3C3-binding and for autophagy.1 Publication
The final 80 residues in the C-terminus define a minimum required region for autophagosome binding called BATS.By similarity
The N-terminal cysteine repeats are required for proper localization to the endoplasmic reticulum.1 Publication

Sequence similaritiesi

Belongs to the ATG14 family.Curated

Keywords - Domaini

Coiled coil

Phylogenomic databases

eggNOGiNOG236057.
GeneTreeiENSGT00390000011854.
HOVERGENiHBG081828.
InParanoidiQ6ZNE5.
KOiK17889.
OMAiCVQSGDF.
OrthoDBiEOG7SV0WR.
PhylomeDBiQ6ZNE5.
TreeFamiTF323392.

Family and domain databases

InterProiIPR018791. UV_resistance/autophagy_Atg14.
[Graphical view]
PfamiPF10186. Atg14. 1 hit.
[Graphical view]

Sequences (2)i

Sequence statusi: Complete.

This entry describes 2 isoformsi produced by alternative splicing. Align

Isoform 1 (identifier: Q6ZNE5-1) [UniParc]FASTAAdd to Basket

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

« Hide

        10         20         30         40         50
MASPSGKGAR ALEAPGCGPR PLARDLVDSV DDAEGLYVAV ERCPLCNTTR
60 70 80 90 100
RRLTCAKCVQ SGDFVYFDGR DRERFIDKKE RLSRLKSKQE EFQKEVLKAM
110 120 130 140 150
EGKWITDQLR WKIMSCKMRI EQLKQTICKG NEEMEKNSEG LLKTKEKNQK
160 170 180 190 200
LYSRAQRHQE KKEKIQRHNR KLGDLVEKKT IDLRSHYERL ANLRRSHILE
210 220 230 240 250
LTSVIFPIEE VKTGVRDPAD VSSESDSAMT SSTVSKLAEA RRTTYLSGRW
260 270 280 290 300
VCDDHNGDTS ISITGPWISL PNNGDYSAYY SWVEEKKTTQ GPDMEQSNPA
310 320 330 340 350
YTISAALCYA TQLVNILSHI LDVNLPKKLC NSEFCGENLS KQKFTRAVKK
360 370 380 390 400
LNANILYLCF SQHVNLDQLQ PLHTLRNLMY LVSPSSEHLG RSGPFEVRAD
410 420 430 440 450
LEESMEFVDP GVAGESDESG DERVSDEETD LGTDWENLPS PRFCDIPSQS
460 470 480 490
VEVSQSQSTQ ASPPIASSSA GGMISSAAAS VTSWFKAYTG HR
Length:492
Mass (Da):55,309
Last modified:July 5, 2004 - v1
Checksum:iA3EAB0580077D7A6
GO
Isoform 2 (identifier: Q6ZNE5-2) [UniParc]FASTAAdd to Basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-113: Missing.

Show »
Length:379
Mass (Da):42,434
Checksum:iD6051FFD708A9943
GO

Sequence cautioni

The sequence BAA74854.2 differs from that shown. Reason: Erroneous initiation. Translation N-terminally shortened.

Experimental Info

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Sequence conflicti256 – 2561N → S in AAI09090. (PubMed:15489334)Curated

Natural variant

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Natural varianti59 – 591V → I.
Corresponds to variant rs57295720 [ dbSNP | Ensembl ].
VAR_061240
Natural varianti131 – 1311N → K.
Corresponds to variant rs17675076 [ dbSNP | Ensembl ].
VAR_049514

Alternative sequence

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Alternative sequencei1 – 113113Missing in isoform 2. 1 PublicationVSP_013931Add
BLAST

Sequence databases

Select the link destinations:
EMBL
GenBank
DDBJ
Links Updated
AB020638 mRNA. Translation: BAA74854.2. Different initiation.
AK131251 mRNA. Translation: BAD18430.1.
AK292810 mRNA. Translation: BAF85499.1.
AL158801 Genomic DNA. No translation available.
CH471061 Genomic DNA. Translation: EAW80673.1.
BC109089 mRNA. Translation: AAI09090.1.
BC109090 mRNA. Translation: AAI09091.1.
BR000826 mRNA. Translation: FAA00433.1.
CCDSiCCDS32087.1. [Q6ZNE5-1]
RefSeqiNP_055739.2. NM_014924.4. [Q6ZNE5-1]
UniGeneiHs.414809.

Genome annotation databases

EnsembliENST00000247178; ENSP00000247178; ENSG00000126775. [Q6ZNE5-1]
GeneIDi22863.
KEGGihsa:22863.
UCSCiuc001xbw.2. human. [Q6ZNE5-1]

Polymorphism databases

DMDMi67461020.

Keywords - Coding sequence diversityi

Alternative splicing, Polymorphism

Cross-referencesi

Sequence databases

Select the link destinations:
EMBL
GenBank
DDBJ
Links Updated
AB020638 mRNA. Translation: BAA74854.2 . Different initiation.
AK131251 mRNA. Translation: BAD18430.1 .
AK292810 mRNA. Translation: BAF85499.1 .
AL158801 Genomic DNA. No translation available.
CH471061 Genomic DNA. Translation: EAW80673.1 .
BC109089 mRNA. Translation: AAI09090.1 .
BC109090 mRNA. Translation: AAI09091.1 .
BR000826 mRNA. Translation: FAA00433.1 .
CCDSi CCDS32087.1. [Q6ZNE5-1 ]
RefSeqi NP_055739.2. NM_014924.4. [Q6ZNE5-1 ]
UniGenei Hs.414809.

3D structure databases

ProteinModelPortali Q6ZNE5.
ModBasei Search...
MobiDBi Search...

Protein-protein interaction databases

BioGridi 116531. 13 interactions.
DIPi DIP-48651N.
IntActi Q6ZNE5. 7 interactions.
MINTi MINT-8304673.

PTM databases

PhosphoSitei Q6ZNE5.

Polymorphism databases

DMDMi 67461020.

Proteomic databases

MaxQBi Q6ZNE5.
PaxDbi Q6ZNE5.
PRIDEi Q6ZNE5.

Protocols and materials databases

Structural Biology Knowledgebase Search...

Genome annotation databases

Ensembli ENST00000247178 ; ENSP00000247178 ; ENSG00000126775 . [Q6ZNE5-1 ]
GeneIDi 22863.
KEGGi hsa:22863.
UCSCi uc001xbw.2. human. [Q6ZNE5-1 ]

Organism-specific databases

CTDi 22863.
GeneCardsi GC14M055834.
HGNCi HGNC:19962. ATG14.
HPAi CAB037242.
HPA053515.
MIMi 613515. gene.
neXtProti NX_Q6ZNE5.
PharmGKBi PA165478560.
HUGEi Search...
GenAtlasi Search...

Phylogenomic databases

eggNOGi NOG236057.
GeneTreei ENSGT00390000011854.
HOVERGENi HBG081828.
InParanoidi Q6ZNE5.
KOi K17889.
OMAi CVQSGDF.
OrthoDBi EOG7SV0WR.
PhylomeDBi Q6ZNE5.
TreeFami TF323392.

Miscellaneous databases

GenomeRNAii 22863.
NextBioi 43375.
PROi Q6ZNE5.
SOURCEi Search...

Gene expression databases

Bgeei Q6ZNE5.
CleanExi HS_KIAA0831.
Genevestigatori Q6ZNE5.

Family and domain databases

InterProi IPR018791. UV_resistance/autophagy_Atg14.
[Graphical view ]
Pfami PF10186. Atg14. 1 hit.
[Graphical view ]
ProtoNeti Search...

Publicationsi

« Hide 'large scale' publications
  1. "Prediction of the coding sequences of unidentified human genes. XII. The complete sequences of 100 new cDNA clones from brain which code for large proteins in vitro."
    Nagase T., Ishikawa K., Suyama M., Kikuno R., Hirosawa M., Miyajima N., Tanaka A., Kotani H., Nomura N., Ohara O.
    DNA Res. 5:355-364(1998) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 2).
    Tissue: Brain.
  2. "Complete sequencing and characterization of 21,243 full-length human cDNAs."
    Ota T., Suzuki Y., Nishikawa T., Otsuki T., Sugiyama T., Irie R., Wakamatsu A., Hayashi K., Sato H., Nagai K., Kimura K., Makita H., Sekine M., Obayashi M., Nishi T., Shibahara T., Tanaka T., Ishii S.
    , Yamamoto J., Saito K., Kawai Y., Isono Y., Nakamura Y., Nagahari K., Murakami K., Yasuda T., Iwayanagi T., Wagatsuma M., Shiratori A., Sudo H., Hosoiri T., Kaku Y., Kodaira H., Kondo H., Sugawara M., Takahashi M., Kanda K., Yokoi T., Furuya T., Kikkawa E., Omura Y., Abe K., Kamihara K., Katsuta N., Sato K., Tanikawa M., Yamazaki M., Ninomiya K., Ishibashi T., Yamashita H., Murakawa K., Fujimori K., Tanai H., Kimata M., Watanabe M., Hiraoka S., Chiba Y., Ishida S., Ono Y., Takiguchi S., Watanabe S., Yosida M., Hotuta T., Kusano J., Kanehori K., Takahashi-Fujii A., Hara H., Tanase T.-O., Nomura Y., Togiya S., Komai F., Hara R., Takeuchi K., Arita M., Imose N., Musashino K., Yuuki H., Oshima A., Sasaki N., Aotsuka S., Yoshikawa Y., Matsunawa H., Ichihara T., Shiohata N., Sano S., Moriya S., Momiyama H., Satoh N., Takami S., Terashima Y., Suzuki O., Nakagawa S., Senoh A., Mizoguchi H., Goto Y., Shimizu F., Wakebe H., Hishigaki H., Watanabe T., Sugiyama A., Takemoto M., Kawakami B., Yamazaki M., Watanabe K., Kumagai A., Itakura S., Fukuzumi Y., Fujimori Y., Komiyama M., Tashiro H., Tanigami A., Fujiwara T., Ono T., Yamada K., Fujii Y., Ozaki K., Hirao M., Ohmori Y., Kawabata A., Hikiji T., Kobatake N., Inagaki H., Ikema Y., Okamoto S., Okitani R., Kawakami T., Noguchi S., Itoh T., Shigeta K., Senba T., Matsumura K., Nakajima Y., Mizuno T., Morinaga M., Sasaki M., Togashi T., Oyama M., Hata H., Watanabe M., Komatsu T., Mizushima-Sugano J., Satoh T., Shirai Y., Takahashi Y., Nakagawa K., Okumura K., Nagase T., Nomura N., Kikuchi H., Masuho Y., Yamashita R., Nakai K., Yada T., Nakamura Y., Ohara O., Isogai T., Sugano S.
    Nat. Genet. 36:40-45(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 1).
    Tissue: Hippocampus and Trachea.
  3. "The DNA sequence and analysis of human chromosome 14."
    Heilig R., Eckenberg R., Petit J.-L., Fonknechten N., Da Silva C., Cattolico L., Levy M., Barbe V., De Berardinis V., Ureta-Vidal A., Pelletier E., Vico V., Anthouard V., Rowen L., Madan A., Qin S., Sun H., Du H.
    , Pepin K., Artiguenave F., Robert C., Cruaud C., Bruels T., Jaillon O., Friedlander L., Samson G., Brottier P., Cure S., Segurens B., Aniere F., Samain S., Crespeau H., Abbasi N., Aiach N., Boscus D., Dickhoff R., Dors M., Dubois I., Friedman C., Gouyvenoux M., James R., Madan A., Mairey-Estrada B., Mangenot S., Martins N., Menard M., Oztas S., Ratcliffe A., Shaffer T., Trask B., Vacherie B., Bellemere C., Belser C., Besnard-Gonnet M., Bartol-Mavel D., Boutard M., Briez-Silla S., Combette S., Dufosse-Laurent V., Ferron C., Lechaplais C., Louesse C., Muselet D., Magdelenat G., Pateau E., Petit E., Sirvain-Trukniewicz P., Trybou A., Vega-Czarny N., Bataille E., Bluet E., Bordelais I., Dubois M., Dumont C., Guerin T., Haffray S., Hammadi R., Muanga J., Pellouin V., Robert D., Wunderle E., Gauguet G., Roy A., Sainte-Marthe L., Verdier J., Verdier-Discala C., Hillier L.W., Fulton L., McPherson J., Matsuda F., Wilson R., Scarpelli C., Gyapay G., Wincker P., Saurin W., Quetier F., Waterston R., Hood L., Weissenbach J.
    Nature 421:601-607(2003) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
  4. Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
  5. "The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)."
    The MGC Project Team
    Genome Res. 14:2121-2127(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 1).
  6. "Beclin 1 forms two distinct phosphatidylinositol 3-kinase complexes with mammalian Atg14 and UVRAG."
    Itakura E., Kishi C., Inoue K., Mizushima N.
    Mol. Biol. Cell 19:5360-5372(2008) [PubMed] [Europe PMC] [Abstract]
    Cited for: IDENTIFICATION, FUNCTION, INTERACTION WITH BECN1; PIK3C3 AND PIK3R4, SUBCELLULAR LOCATION.
  7. "Global, in vivo, and site-specific phosphorylation dynamics in signaling networks."
    Olsen J.V., Blagoev B., Gnad F., Macek B., Kumar C., Mortensen P., Mann M.
    Cell 127:635-648(2006) [PubMed] [Europe PMC] [Abstract]
    Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-29, IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
    Tissue: Cervix carcinoma.
  8. "Identification of Barkor as a mammalian autophagy-specific factor for Beclin 1 and class III phosphatidylinositol 3-kinase."
    Sun Q., Fan W., Chen K., Ding X., Chen S., Zhong Q.
    Proc. Natl. Acad. Sci. U.S.A. 105:19211-19216(2008) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, INTERACTION WITH BECN1, SUBCELLULAR LOCATION.
  9. Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-29; SER-416 AND THR-429, IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
  10. "Two Beclin 1-binding proteins, Atg14L and Rubicon, reciprocally regulate autophagy at different stages."
    Matsunaga K., Saitoh T., Tabata K., Omori H., Satoh T., Kurotori N., Maejima I., Shirahama-Noda K., Ichimura T., Isobe T., Akira S., Noda T., Yoshimori T.
    Nat. Cell Biol. 11:385-396(2009) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, INTERACTION WITH BECN1; PIK3C3 AND PIK3R4, SUBCELLULAR LOCATION.
  11. "Autophagy requires endoplasmic reticulum targeting of the PI3-kinase complex via Atg14L."
    Matsunaga K., Morita E., Saitoh T., Akira S., Ktistakis N.T., Izumi T., Noda T., Yoshimori T.
    J. Cell Biol. 190:511-521(2010) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, SUBCELLULAR LOCATION, DOMAIN, MUTAGENESIS OF CYS-43; CYS-46; CYS-55 AND CYS-58.
  12. "Imperfect interface of Beclin1 coiled-coil domain regulates homodimer and heterodimer formation with Atg14L and UVRAG."
    Li X., He L., Che K.H., Funderburk S.F., Pan L., Pan N., Zhang M., Yue Z., Zhao Y.
    Nat. Commun. 3:662-662(2012) [PubMed] [Europe PMC] [Abstract]
    Cited for: INTERACTION WITH BECN1, SUBCELLULAR LOCATION.
  13. "Beclin 2 functions in autophagy, degradation of G protein-coupled receptors, and metabolism."
    He C., Wei Y., Sun K., Li B., Dong X., Zou Z., Liu Y., Kinch L.N., Khan S., Sinha S., Xavier R.J., Grishin N.V., Xiao G., Eskelinen E.L., Scherer P.E., Whistler J.L., Levine B.
    Cell 154:1085-1099(2013) [PubMed] [Europe PMC] [Abstract]
    Cited for: INTERACTION WITH BECN1P1/BECN2.

Entry informationi

Entry nameiBAKOR_HUMAN
AccessioniPrimary (citable) accession number: Q6ZNE5
Secondary accession number(s): A6NJE4
, A8K9U5, B7ZWP5, O94920, Q32MK7, Q32MK8
Entry historyi
Integrated into UniProtKB/Swiss-Prot: June 7, 2005
Last sequence update: July 5, 2004
Last modified: October 29, 2014
This is version 88 of the entry and version 1 of the sequence. [Complete history]
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program
DisclaimerAny medical or genetic information present in this entry is provided for research, educational and informational purposes only. It is not in any way intended to be used as a substitute for professional medical advice, diagnosis, treatment or care.

Miscellaneousi

Keywords - Technical termi

Complete proteome, Reference proteome

Documents

  1. Human chromosome 14
    Human chromosome 14: entries, gene names and cross-references to MIM
  2. Human entries with polymorphisms or disease mutations
    List of human entries with polymorphisms or disease mutations
  3. Human polymorphisms and disease mutations
    Index of human polymorphisms and disease mutations
  4. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
  5. SIMILARITY comments
    Index of protein domains and families

External Data

Dasty 3