4v7r

X-ray diffraction
4Å resolution

Yeast 80S ribosome.

Released:
Source organism: Saccharomyces cerevisiae
Primary publication:
Crystal structure of the eukaryotic ribosome.
Science 330 1203-9 (2010)
PMID: 21109664

Function and Biology Details

Structure analysis Details

Assemblies composition:
hetero 73-mer (preferred)
hetero 66-mer
PDBe Complex ID:
PDB-CPX-126826 (preferred)
Entry contents:
69 distinct polypeptide molecules
4 distinct RNA molecules
Macromolecules (73 distinct):
Small ribosomal subunit protein uS2A Chains: AA, CA
Molecule details ›
Chains: AA, CA
Length: 252 amino acids
Theoretical weight: 28.05 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P32905 (Residues: 1-252; Coverage: 100%)
Gene names: NAB1, NAB1A, RPS0A, YGR214W, YST1
Sequence domains: Ribosomal protein S2
Small ribosomal subunit protein uS5 Chains: AB, CB
Molecule details ›
Chains: AB, CB
Length: 254 amino acids
Theoretical weight: 27.49 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P25443 (Residues: 1-254; Coverage: 100%)
Gene names: G2893, RPS2, RPS4, SUP38, SUP44, YGL123W
Sequence domains:
Small ribosomal subunit protein uS3 Chains: AC, CC
Molecule details ›
Chains: AC, CC
Length: 240 amino acids
Theoretical weight: 26.54 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05750 (Residues: 1-240; Coverage: 100%)
Gene names: N1653, RPS3, SUF14, YNL178W
Sequence domains:
Small ribosomal subunit protein uS7 Chains: AD, CD
Molecule details ›
Chains: AD, CD
Length: 225 amino acids
Theoretical weight: 25.07 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P26783 (Residues: 1-225; Coverage: 100%)
Gene names: J2045, RPS2, RPS5, YJR123W
Sequence domains: Ribosomal protein S7p/S5e
Small ribosomal subunit protein uS4A Chains: AE, CE
Molecule details ›
Chains: AE, CE
Length: 197 amino acids
Theoretical weight: 22.49 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: O13516 (Residues: 1-197; Coverage: 100%)
Gene names: RPS13A, RPS9A, YPL081W, YS11A
Sequence domains:
40S ribosomal protein S11 Chains: AF, CF
Molecule details ›
Chains: AF, CF
Length: 156 amino acids
Theoretical weight: 17.79 KDa
Source organism: Saccharomyces cerevisiae
Small ribosomal subunit protein uS15 Chains: AG, CG
Molecule details ›
Chains: AG, CG
Length: 151 amino acids
Theoretical weight: 17.06 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05756 (Residues: 1-151; Coverage: 100%)
Gene names: D4252, RPS13, RPS13C, YD9609.18, YDR064W
Sequence domains:
Small ribosomal subunit protein uS11A Chains: AH, CH
Molecule details ›
Chains: AH, CH
Length: 137 amino acids
Theoretical weight: 14.56 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P06367 (Residues: 1-137; Coverage: 100%)
Gene names: CRY1, RPL59, RPS14A, YCR031C, YCR31C
Sequence domains: Ribosomal protein S11
Small ribosomal subunit protein uS19 Chains: AI, CI
Molecule details ›
Chains: AI, CI
Length: 142 amino acids
Theoretical weight: 16.03 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: Q01855 (Residues: 1-142; Coverage: 100%)
Gene names: RPS15, RPS21, YOL040C
Sequence domains: Ribosomal protein S19
40S ribosomal protein S16 Chains: AJ, CJ
Molecule details ›
Chains: AJ, CJ
Length: 143 amino acids
Theoretical weight: 15.88 KDa
Source organism: Saccharomyces cerevisiae
Small ribosomal subunit protein eS17A Chains: AK, CK
Molecule details ›
Chains: AK, CK
Length: 136 amino acids
Theoretical weight: 15.82 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P02407 (Residues: 1-136; Coverage: 100%)
Gene names: RP51A, RPS17A, YML024W
Sequence domains: Ribosomal S17
40S ribosomal protein S18 Chains: AL, CL
Molecule details ›
Chains: AL, CL
Length: 146 amino acids
Theoretical weight: 17.07 KDa
Source organism: Saccharomyces cerevisiae
Small ribosomal subunit protein eS19A Chains: AM, CM
Molecule details ›
Chains: AM, CM
Length: 144 amino acids
Theoretical weight: 15.94 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P07280 (Residues: 1-144; Coverage: 100%)
Gene names: RP55A, RPS16AA, RPS19A, YOL121C
Sequence domains: Ribosomal protein S19e
Small ribosomal subunit protein uS10 Chains: AN, CN
Molecule details ›
Chains: AN, CN
Length: 121 amino acids
Theoretical weight: 13.93 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P38701 (Residues: 1-121; Coverage: 100%)
Gene names: RPS20, URP2, YHL015W
Sequence domains: Ribosomal protein S10p/S20e
Small ribosomal subunit protein uS8A Chains: AO, CO
Molecule details ›
Chains: AO, CO
Length: 130 amino acids
Theoretical weight: 14.65 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P0C0W1 (Residues: 1-130; Coverage: 100%)
Gene names: J0355, RPS22A, RPS24, RPS24A, YJL190C
Sequence domains: Ribosomal protein S8
40S ribosomal protein S23 Chains: AP, CP
Molecule details ›
Chains: AP, CP
Length: 145 amino acids
Theoretical weight: 16.07 KDa
Source organism: Saccharomyces cerevisiae
Small ribosomal subunit protein eS25A Chains: AQ, CQ
Molecule details ›
Chains: AQ, CQ
Length: 108 amino acids
Theoretical weight: 12.07 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: Q3E792 (Residues: 1-108; Coverage: 100%)
Gene names: RPS25A, RPS31A, YGR027C
Sequence domains: S25 ribosomal protein
Small ribosomal subunit protein eS28A Chains: AR, CR
Molecule details ›
Chains: AR, CR
Length: 67 amino acids
Theoretical weight: 7.61 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: Q3E7X9 (Residues: 1-67; Coverage: 100%)
Gene names: O3600, RPS28A, RPS33, RPS33A, YOR167C
Sequence domains: Ribosomal protein S28e
Small ribosomal subunit protein uS14A Chains: AS, CS
Molecule details ›
Chains: AS, CS
Length: 56 amino acids
Theoretical weight: 6.68 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P41057 (Residues: 1-56; Coverage: 100%)
Gene names: L8084.11, RPS29A, RPS36A, YLR388W, YS29A
Sequence domains: Ribosomal protein S14p/S29e
Small ribosomal subunit protein RACK1 Chains: AT, CT
Molecule details ›
Chains: AT, CT
Length: 319 amino acids
Theoretical weight: 34.84 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P38011 (Residues: 1-319; Coverage: 100%)
Gene names: ASC1, CPC2, YM9718.15C, YMR116C
Sequence domains: WD domain, G-beta repeat
Unassigned secondary structure Chains: Aa, Bo, Ca
Molecule details ›
Chains: Aa, Bo, Ca
Length: 20 amino acids
Theoretical weight: 1.72 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Ab, Cb
Molecule details ›
Chains: Ab, Cb
Length: 105 amino acids
Theoretical weight: 8.95 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Ac, Cc
Molecule details ›
Chains: Ac, Cc
Length: 93 amino acids
Theoretical weight: 7.93 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Ad, Cd
Molecule details ›
Chains: Ad, Cd
Length: 35 amino acids
Theoretical weight: 3 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Ae, Bj, Dj
Molecule details ›
Chains: Ae, Bj, Dj
Length: 21 amino acids
Theoretical weight: 1.81 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chain: Af
Molecule details ›
Chain: Af
Length: 11 amino acids
Theoretical weight: 954 Da
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Ah, Ch
Molecule details ›
Chains: Ah, Ch
Length: 41 amino acids
Theoretical weight: 3.51 KDa
Source organism: Saccharomyces cerevisiae
60S ribosomal protein L1 Chains: BA, DA
Molecule details ›
Chains: BA, DA
Length: 217 amino acids
Theoretical weight: 24.52 KDa
Source organism: Saccharomyces cerevisiae
60S ribosomal protein L2 Chains: BB, DB
Molecule details ›
Chains: BB, DB
Length: 254 amino acids
Theoretical weight: 27.46 KDa
Source organism: Saccharomyces cerevisiae
Large ribosomal subunit protein uL3 Chains: BC, DC
Molecule details ›
Chains: BC, DC
Length: 387 amino acids
Theoretical weight: 43.85 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P14126 (Residues: 1-387; Coverage: 100%)
Gene names: MAK8, RPL3, TCM1, YOR063W, YOR29-14
Sequence domains: Ribosomal protein L3
Large ribosomal subunit protein uL4A Chains: BD, DD
Molecule details ›
Chains: BD, DD
Length: 362 amino acids
Theoretical weight: 39.16 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P10664 (Residues: 1-362; Coverage: 100%)
Gene names: RPL2, RPL2A, RPL4A, YBR0315, YBR031W
Sequence domains:
Large ribosomal subunit protein uL18 Chains: BE, DE
Molecule details ›
Chains: BE, DE
Length: 297 amino acids
Theoretical weight: 33.79 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P26321 (Residues: 1-297; Coverage: 100%)
Gene names: LPI14W, RPL1, RPL1A, RPL5, YPL131W
Sequence domains:
Large ribosomal subunit protein eL6A Chains: BF, DF
Molecule details ›
Chains: BF, DF
Length: 176 amino acids
Theoretical weight: 20 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: Q02326 (Residues: 1-176; Coverage: 100%)
Gene names: RPL17A, RPL6A, YL16A, YML073C
Sequence domains: Ribosomal protein L6e
Large ribosomal subunit protein uL30A Chains: BG, DG
Molecule details ›
Chains: BG, DG
Length: 244 amino acids
Theoretical weight: 27.69 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05737 (Residues: 1-244; Coverage: 100%)
Gene names: RPL6A, RPL7A, RPL8A, YGL076C, YL8A
Sequence domains:
Large ribosomal subunit protein eL8A Chains: BH, DH
Molecule details ›
Chains: BH, DH
Length: 256 amino acids
Theoretical weight: 28.18 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P17076 (Residues: 1-256; Coverage: 100%)
Gene names: MAK7, RPL4A, RPL8A, YHL033C
Sequence domains: Ribosomal protein L7Ae/L30e/S12e/Gadd45 family
Large ribosomal subunit protein uL6A Chains: BI, DI
Molecule details ›
Chains: BI, DI
Length: 191 amino acids
Theoretical weight: 21.61 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05738 (Residues: 1-191; Coverage: 100%)
Gene names: RPL8A, RPL9, RPL9A, YGL147C
Sequence domains: Ribosomal protein L6
Large ribosomal subunit protein uL16 Chains: BJ, DJ
Molecule details ›
Chains: BJ, DJ
Length: 221 amino acids
Theoretical weight: 25.41 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P41805 (Residues: 1-221; Coverage: 100%)
Gene names: GRC5, QSR1, RPL10, YLR075W
Sequence domains: Ribosomal protein L16p/L10e
Large ribosomal subunit protein uL5A Chains: BK, DK
Molecule details ›
Chains: BK, DK
Length: 174 amino acids
Theoretical weight: 19.76 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P0C0W9 (Residues: 1-174; Coverage: 100%)
Gene names: P8283.14, RP39A, RPL11A, RPL16A, YPR102C
Sequence domains:
Large ribosomal subunit protein eL14A Chains: BN, DN
Molecule details ›
Chains: BN, DN
Length: 138 amino acids
Theoretical weight: 15.2 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P36105 (Residues: 1-138; Coverage: 100%)
Gene names: RPL14A, YKL006W, YKL153
Sequence domains:
Large ribosomal subunit protein eL15A Chains: BO, DO
Molecule details ›
Chains: BO, DO
Length: 204 amino acids
Theoretical weight: 24.48 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05748 (Residues: 1-204; Coverage: 100%)
Gene names: RPL10A, RPL13A, RPL15A, YL10A, YLR029C
Sequence domains: Ribosomal L15
Large ribosomal subunit protein uL13A Chains: BP, DP
Molecule details ›
Chains: BP, DP
Length: 199 amino acids
Theoretical weight: 22.25 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P26784 (Residues: 1-199; Coverage: 100%)
Gene names: RPL13, RPL16A, RPL21A, YIL133C
Sequence domains: Ribosomal protein L13
Large ribosomal subunit protein uL22A Chains: BQ, DQ
Molecule details ›
Chains: BQ, DQ
Length: 184 amino acids
Theoretical weight: 20.59 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05740 (Residues: 1-184; Coverage: 100%)
Gene names: RPL17, RPL17A, RPL20A, YKL180W
Sequence domains: Ribosomal protein L22p/L17e
60S ribosomal protein L18 Chains: BR, DR
Molecule details ›
Chains: BR, DR
Length: 186 amino acids
Theoretical weight: 20.61 KDa
Source organism: Saccharomyces cerevisiae
60S ribosomal protein L19 Chains: BS, DS
Molecule details ›
Chains: BS, DS
Length: 189 amino acids
Theoretical weight: 21.76 KDa
Source organism: Saccharomyces cerevisiae
Large ribosomal subunit protein eL21A Chains: BT, DT
Molecule details ›
Chains: BT, DT
Length: 160 amino acids
Theoretical weight: 18.28 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: Q02753 (Residues: 1-160; Coverage: 100%)
Gene names: RPL21A, URP1, YBR1401, YBR191W
Sequence domains: Ribosomal protein L21e
60S ribosomal protein L23 Chains: BU, DU
Molecule details ›
Chains: BU, DU
Length: 137 amino acids
Theoretical weight: 14.49 KDa
Source organism: Saccharomyces cerevisiae
Large ribosomal subunit protein eL24A Chains: BV, DV
Molecule details ›
Chains: BV, DV
Length: 155 amino acids
Theoretical weight: 17.66 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P04449 (Residues: 1-155; Coverage: 100%)
Gene names: RP29, RPL24A, RPL30A, YGL031C
Sequence domains: Ribosomal protein L24e
Large ribosomal subunit protein uL23 Chains: BW, DW
Molecule details ›
Chains: BW, DW
Length: 142 amino acids
Theoretical weight: 15.79 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P04456 (Residues: 1-142; Coverage: 100%)
Gene names: RPL25, YOL127W
Sequence domains:
Large ribosomal subunit protein uL24A Chains: BX, DX
Molecule details ›
Chains: BX, DX
Length: 127 amino acids
Theoretical weight: 14.27 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05743 (Residues: 1-127; Coverage: 100%)
Gene names: L8300.4, RPL26, RPL26A, RPL33A, YLR344W
Sequence domains:
Large ribosomal subunit protein uL15 Chains: BY, DY
Molecule details ›
Chains: BY, DY
Length: 149 amino acids
Theoretical weight: 16.76 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P02406 (Residues: 1-149; Coverage: 100%)
Gene names: CYH2, RPL28, YGL103W
Sequence domains: Ribosomal proteins 50S-L15, 50S-L18e, 60S-L27A
Large ribosomal subunit protein eL30 Chains: BZ, DZ
Molecule details ›
Chains: BZ, DZ
Length: 105 amino acids
Theoretical weight: 11.43 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P14120 (Residues: 1-105; Coverage: 100%)
Gene names: RPL30, RPL32, YGL030W
Sequence domains: Ribosomal protein L7Ae/L30e/S12e/Gadd45 family
Large ribosomal subunit protein eL31A Chains: Ba, Da
Molecule details ›
Chains: Ba, Da
Length: 113 amino acids
Theoretical weight: 12.98 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P0C2H8 (Residues: 1-113; Coverage: 100%)
Gene names: D2478, RPL31A, RPL34, RPL34A, YDL075W
Sequence domains: Ribosomal protein L31e
Large ribosomal subunit protein eL32 Chains: Bb, Db
Molecule details ›
Chains: Bb, Db
Length: 130 amino acids
Theoretical weight: 14.81 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P38061 (Residues: 1-130; Coverage: 100%)
Gene names: RPL32, YBL0838, YBL092W
Sequence domains: Ribosomal protein L32
60S ribosomal protein L35 Chains: Bc, Dc
Molecule details ›
Chains: Bc, Dc
Length: 120 amino acids
Theoretical weight: 13.94 KDa
Source organism: Saccharomyces cerevisiae
Large ribosomal subunit protein eL37A Chains: Bd, Dd
Molecule details ›
Chains: Bd, Dd
Length: 88 amino acids
Theoretical weight: 9.88 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P49166 (Residues: 1-88; Coverage: 100%)
Gene names: L9470.6, RPL35A, RPL37A, YLR185W
Sequence domains: Ribosomal protein L37e
Large ribosomal subunit protein eL39 Chains: Be, De
Molecule details ›
Chains: Be, De
Length: 51 amino acids
Theoretical weight: 6.36 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P04650 (Residues: 1-51; Coverage: 100%)
Gene names: J0360, RPL39, RPL46, SPB2, YJL189W
Sequence domains: Ribosomal L39 protein
60S ribosomal protein L42 Chains: Bf, Df
Molecule details ›
Chains: Bf, Df
Length: 106 amino acids
Theoretical weight: 12.25 KDa
Source organism: Saccharomyces cerevisiae
60S ribosomal protein L43 Chains: Bg, Dg
Molecule details ›
Chains: Bg, Dg
Length: 92 amino acids
Theoretical weight: 10.11 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Bh, Dh
Molecule details ›
Chains: Bh, Dh
Length: 44 amino acids
Theoretical weight: 3.76 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Bi, Di
Molecule details ›
Chains: Bi, Di
Length: 12 amino acids
Theoretical weight: 1.04 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chains: Bk, Dk
Molecule details ›
Chains: Bk, Dk
Length: 16 amino acids
Theoretical weight: 1.38 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chain: Bl
Molecule details ›
Chain: Bl
Length: 19 amino acids
Theoretical weight: 1.64 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chain: Bm
Molecule details ›
Chain: Bm
Length: 9 amino acids
Theoretical weight: 784 Da
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chain: Bn
Molecule details ›
Chain: Bn
Length: 27 amino acids
Theoretical weight: 2.32 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chain: Bp
Molecule details ›
Chain: Bp
Length: 8 amino acids
Theoretical weight: 699 Da
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chain: Bq
Molecule details ›
Chain: Bq
Length: 17 amino acids
Theoretical weight: 1.46 KDa
Source organism: Saccharomyces cerevisiae
Unassigned secondary structure Chain: Br
Molecule details ›
Chain: Br
Length: 23 amino acids
Theoretical weight: 1.98 KDa
Source organism: Saccharomyces cerevisiae
60S ribosomal protein L12 Chain: DL
Molecule details ›
Chain: DL
Length: 165 amino acids
Theoretical weight: 17.85 KDa
Source organism: Saccharomyces cerevisiae
Large ribosomal subunit protein uL10 Chain: DM
Molecule details ›
Chain: DM
Length: 312 amino acids
Theoretical weight: 33.8 KDa
Source organism: Saccharomyces cerevisiae
UniProt:
  • Canonical: P05317 (Residues: 1-312; Coverage: 100%)
Gene names: L10E, L8300.8, RPA0, RPL10E, RPLA0, RPP0, YLR340W
Sequence domains:
18S Ribosomal RNA Chains: A1, C1
Molecule details ›
Chains: A1, C1
Length: 1800 nucleotides
Theoretical weight: 579.76 KDa
Sequence domains: Eukaryotic small subunit ribosomal RNA
25S ribosomal RNA Chains: B1, D1
Molecule details ›
Chains: B1, D1
Length: 3396 nucleotides
Theoretical weight: 1097.49 KDa
Sequence domains: Eukaryotic large subunit ribosomal RNA
5S ribosomal RNA Chains: B2, D2
Molecule details ›
Chains: B2, D2
Length: 121 nucleotides
Theoretical weight: 38.95 KDa
Sequence domains: 5S ribosomal RNA
5.8S ribosomal RNA Chains: B3, D3
Molecule details ›
Chains: B3, D3
Length: 158 nucleotides
Theoretical weight: 50.68 KDa
Sequence domains: 5.8S ribosomal RNA

Ligands and Environments

1 bound ligand:
No modified residues

Experiments and Validation Details

Entry percentile scores
X-ray source: SLS BEAMLINE X06SA
Spacegroup: P21
Unit cell:
a: 437.11Å b: 288.38Å c: 306.56Å
α: 90° β: 99.13° γ: 90°
R-values:
R R work R free
0.28 0.278 0.341