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Structure paper

TitleStructural visualization of the molecular evolution of CRISPR-Cas9.
Journal, issue, pagesNat Struct Mol Biol, Vol. 33, Issue 2, Page 304-317, Year 2026
Publish dateJan 30, 2026
AuthorsNaoto Nagahata / Kazuki Kato / Sota Yamada / Soumya Kannan / Sae Okazaki / Yukari Isayama / Masahiro Hiraizumi / Keitaro Yamashita / Eugene V Koonin / Feng Zhang / Hiroshi Nishimasu /
PubMed AbstractRNA-guided DNA nucleases Cas9 and IscB (insertion sequences Cas9-like OrfB) are components of type II CRISPR-Cas adaptive immune systems and transposon-associated OMEGA (obligate mobile element- ...RNA-guided DNA nucleases Cas9 and IscB (insertion sequences Cas9-like OrfB) are components of type II CRISPR-Cas adaptive immune systems and transposon-associated OMEGA (obligate mobile element-guided activity) systems, respectively. Sequence and structural comparisons indicate that IscB (~500 residues) evolved into Cas9 (~700-1,600 residues) through protein expansion coupled with guide RNA miniaturization. However, the specific sequence of events in this evolutionary transition remains unknown. Here, we report cryo-electron microscopy structures of four phylogenetically diverse RNA-guided nucleases-two IscBs and two Cas9s-each in complex with its cognate guide RNA and target DNA. Comparisons of these four complex structures to previously reported IscB and Cas9 structures indicate that evolution from IscB to Cas9 involved the loss of the N-terminal PLMP domain and the acquisition of the zinc-finger-containing REC3 domain, followed by bridge helix extension and REC1 domain acquisition. These structural changes led to expansion of the REC lobe, increasing the target DNA cleavage specificity. Additionally, the structural conservation of the RNA scaffolds indicates that the dual CRISPR RNA (crRNA) and trans-activating crRNA guides of CRISPR-Cas9 evolved from the single ωRNA guides of OMEGA systems. Our findings provide insights into the succession of structural changes involved in the exaptation of transposon-associated RNA-guided nucleases for the role of effector nucleases in adaptive immune systems.
External linksNat Struct Mol Biol / PubMed:41617991
MethodsEM (single particle)
Resolution2.38 - 3.17 Å
Structure data

62001

9k2z

EMDB-62001, PDB-9k2z:
Cryo-EM structure of the HfmIscB-omega RNA-target DNA complex
Method: EM (single particle) / Resolution: 2.78 Å

62002

9k30

EMDB-62002, PDB-9k30:
Cryo-EM structure of the TbaIscB-omega RNA-target DNA complex
Method: EM (single particle) / Resolution: 3.17 Å

62003

9k31

EMDB-62003, PDB-9k31:
Cryo-EM structure of the YnpsCas9-guide RNA-target DNA complex
Method: EM (single particle) / Resolution: 2.38 Å

62004

9k32

EMDB-62004, PDB-9k32:
Cryo-EM structure of the NbaCas9-guide RNA-target DNA complex
Method: EM (single particle) / Resolution: 2.85 Å

Chemicals

ChemComp-MG:
Unknown entry

ChemComp-HOH:
WATER

ChemComp-ZN:
Unknown entry

Source
  • metagenome (others)
  • synthetic construct (others)
  • tissierellia bacterium (bacteria)
  • nitrospirota bacterium (bacteria)
KeywordsRNA BINDING PROTEIN / RNA-guided DNA nuclease

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