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Acknowledgements

SEC-SAXS experiments at Photon Factory were performed under Proposal No. 2017G100

and 2018G680. SEC-iCM-SANS, and iCM-SANS at Institut Laue Langevin were performed

under Proposals No. 8-03-854, 8-03-884, 8-03-924, and 8-03-938. Analytical ultracentrifuge

and SAXS experiment at Institute for Integrated Radiation and Nuclear Science, Kyoto

University (KURNS) were performed under Proposal Numbers, 31006, R2024, and R3063 to

H.Y. and R3003 to M.Y.-U. This work was supported by the JSPS Research Fellowship for

Young Scientists (to Y.Y. and R.Y.) and the Grants-in-Aid for Scientific Research (Grant

Numbers, JP18J21063 to Y.Y., JP18K06101 to A.Mat., JP19K16088 and JP21K15051 to K.M.,

JP17K07816 to N.S., JP20K06579 and JP19KK0071 to R.I., JP20K22629 to M.Sh.,

JP18H05229 and JP18H03681 to M.Su., and JP18H05534 to M.Su. and H.K.) from the

Ministry of Education, Culture, Sports, Science and Technology (MEXT), Japan. The study

was partially supported by the Platform Project for Supporting Drug Discovery and Life

Science Research (Basis for Supporting Innovative Drug Discovery and Life Science Research

(BINDS)) from AMED (JP21am0101106, support number 2121) to A.Mat. and H.K. This

study was also supported by Joint Research of the Exploratory Research Center on Life and

Living Systems (ExCELLS) (ExCELLS program No. 21-408 to K.M., No. 20-326 and 21-315

to M.Su., and No. 20-308 and 21-308 to H.Y.), by Joint Research by Institute for Molecular

Science (IMS program No. 19-205, 20-207 and 21-217 to M.Su.), by Research Fund for

Young Scientists in Kyoto University (to K.M.) and by Fund for Project Research in KURNS

(to Y.Y., K.M., M.Sh., and A.O.). This work was also partially supported by the project for

Construction of the basis for the advanced materials science and analytical study by the

innovative use of quantum beam and nuclear sciences in KURNS. Computations were

carried out at National Institutes for Quantum Science and Technology, at Japan Atomic

Energy Agency, and at Tokyo Institute of Technology Global Scientific Information and

Computing Center.

Author contributions

This research was designed by H.K., H.Y., K.K., and M.Su. Sample preparation was

performed by Y.Y., A.O., R.U., K.T., and H.Y. SEC-SAXS experiment was performed by

R.I., M.Y.-U., M.Sh., and M.Su. SEC-iCM-SANS experiments were performed by A.Mar.,

L.P., N.S., R.Y., T.T., and M.Su. AUC experiment was performed by Y.Y. and K.M.

Computational modeling and molecular dynamics simulation were performed by A.Mat.

and H.K. Integrative analysis was performed by Y.Y., A.Mat., H.K., H.Y., K.K., and M.Su.

All the authors wrote the paper.

Competing interests

The authors declare no competing interests.

Additional information

Supplementary information The online version contains supplementary material

available at https://doi.org/10.1038/s42003-022-03143-z.

Correspondence and requests for materials should be addressed to Hidetoshi Kono,

Hirokazu Yagi, Koichi Kato or Masaaki Sugiyama.

Peer review information Communications Biology thanks Carlo Camilloni and Ralf

Biehl for their contribution to the peer review of this work. Primary Handling Editor:

Gene Chong. Peer reviewer reports are available.

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