9SHZ: Histone H2A

Cryo-EM structure of the catalytic core of human telomerase at the elongation state of the repeat addition cycle. Determined by electron microscopy at 3.2 Å resolution. Released 28 Jan 2026.

Method
Electron microscopy
Resolution
3.2 Å
Organism
Homo sapiens
Chains
6
Atoms
15,611
Mol. weight
363.88 kDa
Ligands
A1ACD
Released
28 Jan 2026

Explore 9SHZ in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9SHZ contains 71 α-helices and 46 β-strands across 4 chains. Residue ranges use author residue numbering, as in the PDB file, from PDBe. To see them in 3D, choose the Cartoon representation with Secondary structure coloring: helices, sheets and coils get different colors.

Chain A: 57 helices, 36 β-strands

ElementResiduesLengthSheet
α-helix8-158
β-strand20-2344
α-helix24-318
α-helix33-375
α-helix45-539
β-strand54-5854
α-helix62-643
α-helix77-9115
α-helix96-983
β-strand101-10335
β-strand119-12135
α-helix126-1316
α-helix135-15521
β-strand157-16154
β-strand167-16934
α-helix175-1773
β-strand325-32736
α-helix328-3314
β-strand33217
α-helix341-3422
α-helix346-3494
α-helix354-3618
α-helix362-3665
α-helix379-3835
α-helix384-3874
α-helix390-40011
α-helix405-4128
α-helix445-4528
β-strand45617
α-helix458-47215
α-helix481-49515
β-strand49818
β-strand502-50436
α-helix505-5095
α-helix518-5203
α-helix531-54717
α-helix548-5536
α-helix554-5607
β-strand561-56667
β-strand569-57797
α-helix578-59619
α-helix5981
β-strand599-60029
α-helix601-6022
α-helix603-6119
α-helix615-6162
β-strand617-626109
β-strand629-63689
α-helix637-6393
α-helix650-6523
α-helix654-67118
α-helix673-6764
β-strand679110
α-helix683-69715
α-helix703-7053
β-strand707-712610
α-helix714-7163
α-helix722-73211
β-strand738-739211
β-strand740-749104
β-strand755-764104
α-helix773-78311
β-strand790-79564
β-strand800-801211
α-helix802-81312
β-strand816-82059
β-strand823-82649
α-helix838-85316
α-helix855-8584
β-strand862-865410
β-strand869-874610
α-helix877-88812
β-strand891112
α-helix892-8943
β-strand896112
α-helix900-9023
β-strand904-905210
α-helix912-9154
β-strand920-921210
β-strand927-930413
β-strand933-936413
β-strand942-944313
β-strand953114
α-helix955-9573
α-helix966-98116
α-helix984-9874
α-helix994-101623
α-helix1025-10273
α-helix1029-104921
α-helix1067-108418
α-helix1086-110621
α-helix1109-111911
β-strand1131114
Chain L: 5 helices, 2 β-strands
ElementResiduesLengthSheet
α-helix20-223
α-helix28-3710
β-strand43-4421
α-helix47-7226
β-strand78-7922
α-helix81-9010
α-helix92-987
Chain M: 4 helices, 2 β-strands
ElementResiduesLengthSheet
α-helix39-4911
β-strand54-5522
α-helix57-8428
β-strand89-9021
α-helix92-10211
α-helix108-12417
Chain O: 5 helices, 6 β-strands
ElementResiduesLengthSheet
α-helix98-1036
β-strand113-122103
β-strand142-14763
β-strand152-15763
α-helix159-1646
α-helix170-1723
β-strand180-190113
β-strand203-215133
β-strand22313
α-helix224-2263
α-helix228-2369

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
Histone H2ALprotein130Homo sapiensB2R5B3 (AlphaFold model)
Histone H2BMprotein166Homo sapiensB4DR52 (AlphaFold model)
DNA (5'-d(p*tp*ap*gp*gp*gp*tp*tp*ap*gp*gp*g)-3')NDNA30Homo sapiens
Adrenocortical dysplasia protein homologOprotein458Homo sapiensQ96AP0 (AlphaFold model)
hTR, human telomerase RNABRNA451Homo sapiens
Telomerase reverse transcriptaseAprotein1132Homo sapiensO14746 (AlphaFold model)
Sequence of entity 1 (L), FASTA
>9SHZ_1 Histone H2A (chains L)
MSGRGKQGGKARAKAKTRSSRAGLQFPVGRVRRLLRKGNYAERVGAGAPVYLAAVLEYLT
AEILELAGNAARDNKKTRIIPRHLQLAIRNDEELNKLLGKVTIAQGGVLPNIQAVLLPKK
TESHHKAKGK
Sequence of entity 2 (M), FASTA
>9SHZ_2 Histone H2B (chains M)
MPDPAKSAPAPKKGSKKAVTKVQKKDGKKRKRSRKESYSVYVYKVLKQVHPDTGISSKAM
GIMNSFVNDIFERIAGEASRLAHYNKRSTITSREIQTAVRLLLPGELAKHAVSEGTKAVT
KYTSSNPRNLSPTKPGGSEDRQPPPSQLSAIPPFCLVLRAGIAGQV
Sequence of entity 3 (N), FASTA
>9SHZ_3 DNA (5'-D(P*TP*AP*GP*GP*GP*TP*TP*AP*GP*GP*G)-3') (chains N)
TTAGGGTTAGGGTTAGGGTTAGGGTTAGGG
Sequence of entity 4 (O), FASTA
>9SHZ_4 Adrenocortical dysplasia protein homolog (chains O)
MAGSGRLVLRPWIRELILGSETPSSPRAGQLLEVLQDAEAAVAGPSHAPDTSDVGATLLV
SDGTHSVRCLVTREALDTSDWEEKEFGFRGTEGRLLLLQDCGVHVQVAEGGAPAEFYLQV
DRFSLLPTEQPRLRVPGCNQDLDVQKKLYDCLEEHLSESTSSNAGLSLSQLLDEMREDQE
HQGALVCLAESCLTLEGPCTAPPVTHWAASRCKATGEAVYTVPSSMLCISENDQLILSSL
GPCQRTQGPELPPPDPALQDLSLTLIASPPSSPSSSGTPALPGHMSSEESGTSISLLPAL
SLAAPDPGQRSSSQPSPAICSAPATLTPRSPHASRTPSSPLQSCTPSLSPRSHVPSPHQA
LVTRPQKPSLEFKEFVGLPCKNRPPFPRTGATRGAQEPCSVWEPPKRHRDGSAFQYEYEP
PCTSLCARVQAVRLPPQLMAWALHFLMDAQPGSEPTPM
Sequence of entity 5 (B), FASTA
>9SHZ_5 hTR, human telomerase RNA (chains B)
GGGUUGCGGAGGGUGGGCCUGGGAGGGGUGGUGGCCAUUUUUUGUCUAACCCUAACUGAG
AAGGGCGUAGGCGCCGUGCUUUUGCUCCCCGCGCGCUGUUUUUCUCGCUGACUUUCAGCG
GGCGGAAAAGCCUCGGCCUGCCGCCUUCCACCGUUCAUUCUAGAGCAAACAAAAAAUGUC
AGCUGCUGGCCCGUUCGCCCCUCCCGGGGACCUGCGGCGGGUCGCCUGCCCAGCCCCCGA
ACCCCGCCUGGAGGCCGCGGUCGGCCCGGGGCUUCUCCGGAGGCACCCACUGCCACCGCG
AAGAGUUGGGCUCUGUCAGCCGCGGGUCUCUCGGGGGCGAGGGCGAGGUUCAGGCCUUUC
AGGCCGCAGGAAGAGGAACGGAGCGAGUCCCCGCGCGCGGCGCGAUUCCCUGAGCUGUGG
GACGUGCACCCAGGACUCGGCUCACACAUGC
Sequence of entity 6 (A), FASTA
>9SHZ_6 Telomerase reverse transcriptase (chains A)
MPRAPRCRAVRSLLRSHYREVLPLATFVRRLGPQGWRLVQRGDPAAFRALVAQCLVCVPW
DARPPPAAPSFRQVSCLKELVARVLQRLCERGAKNVLAFGFALLDGARGGPPEAFTTSVR
SYLPNTVTDALRGSGAWGLLLRRVGDDVLVHLLARCALFVLVAPSCAYQVCGPPLYQLGA
ATQARPPPHASGPRRRLGCERAWNHSVREAGVPLGLPAPGARRRGGSASRSLPLPKRPRR
GAAPEPERTPVGQGSWAHPGRTRGPSDRGFCVVSPARPAEEATSLEGALSGTRHSHPSVG
RQHHAGPPSTSRPPRPWDTPCPPVYAETKHFLYSSGDKEQLRPSFLLSSLRPSLTGARRL
VETIFLGSRPWMPGTPRRLPRLPQRYWQMRPLFLELLGNHAQCPYGVLLKTHCPLRAAVT
PAAGVCAREKPQGSVAAPEEEDTDPRRLVQLLRQHSSPWQVYGFVRACLRRLVPPGLWGS
RHNERRFLRNTKKFISLGKHAKLSLQELTWKMSVRDCAWLRRSPGVGCVPAAEHRLREEI
LAKFLHWLMSVYVVELLRSFFYVTETTFQKNRLFFYRKSVWSKLQSIGIRQHLKRVQLRE
LSEAEVRQHREARPALLTSRLRFIPKPDGLRPIVNMDYVVGARTFRREKRAERLTSRVKA
LFSVLNYERARRPGLLGASVLGLDDIHRAWRTFVLRVRAQDPPPELYFVKVDVTGAYDTI
PQDRLTEVIASIIKPQNTYCVRRYAVVQKAAHGHVRKAFKSHVSTLTDLQPYMRQFVAHL
QETSPLRDAVVIEQSSSLNEASSGLFDVFLRFMCHHAVRIRGKSYVQCQGIPQGSILSTL
LCSLCYGDMENKLFAGIRRDGLLLRLVDDFLLVTPHLTHAKTFLRTLVRGVPEYGCVVNL
RKTVVNFPVEDEALGGTAFVQMPAHGLFPWCGLLLDTRTLEVQSDYSSYARTSIRASLTF
NRGFKAGRNMRRKLFGVLRLKCHSLFLDLQVNSLQTVCTNIYKILLLQAYRFHACVLQLP
FHQQVWKNPTFFLRVISDTASLCYSILKAKNAGMSLGAKGAAGPLPSEAVQWLCHQAFLL
KLTRHRVTYVPLLGSLRTAQTQLSRKLPGTTLTALEAAANPALPSDFKTILD

Ligands and cofactors

IDNameFormulaCopies
A1ACD1-{2-deoxy-5-O-[(R)-hydroxy{[(S)-hydroxy(phosphonooxy)phosphoryl]methyl}phospho…C11 H19 N2 O13 P31

Primary citation

Structures of nucleotide-bound human telomerase at several steps of its telomeric DNA repeat addition cycle. Balch, S., Franco-Echevarria, E., Ghanim, G.E. et al. Nat Commun (2026) 17:1847-1847. DOI 10.1038/s41467-026-68560-8 · PubMed

Other PDB entries of the same protein (UniProt B2R5B3 (AlphaFold model), which also has an AlphaFold model), best resolution first:

Browse structure collections

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