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Type 51 R-body assembles from four Reb proteins and undergoes rapid, repeatable \u223c50-fold extension. Although R-bodies were described more than 70 years ago, the roles of RebC and RebD remained unclear, whereas RebA and RebB are proposed as major components. Here, we characterize the wild-type R-body (Rb_WT) and reb gene knockout mutants using integrated biophysical analyses. RebD was incorporated as a minor component, whereas RebC was not detectably incorporated into the final assembly. Mutants lacking RebA or RebB formed rolls but lost pH-dependent extension and the ordered lamellar architecture characteristic of Rb_WT. In contrast, mutants lacking RebC and\/or RebD exhibited reduced roll formation and formed \u03b2-sheet-rich aggregation. Thus, pH-responsive actuation is dictated not simply by major architectural components alone, but by a regulated assembly pathway that builds the ordered lamellar architecture.<\/jats:p>","DOI":"10.1021\/acs.biomac.6c01332","type":"journal-article","created":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T20:01:43Z","timestamp":1789761703000},"page":"6728-6737","source":"Crossref","is-referenced-by-count":0,"title":["Assembly-Pathway Regulation Dictates pH-Responsive Actuation in the R-Body Protein Machinery"],"prefix":"10.1021","volume":"27","author":[{"given":"Koki","family":"Date","sequence":"first","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2998-9049","authenticated-orcid":true,"given":"Kosuke","family":"Kikuchi","sequence":"additional","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tatsuya","family":"Niwa","sequence":"additional","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]},{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0878-6488","authenticated-orcid":true,"given":"Hironari","family":"Kamikubo","sequence":"additional","affiliation":[{"name":"Nara Institute of Science and Technology (NAIST) , , 8916-5 Takayama-cho , ,","place":["Ikoma, Japan, 630-0192"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sota","family":"Masumura","sequence":"additional","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3120-3789","authenticated-orcid":true,"given":"Thuc Toan","family":"Pham","sequence":"additional","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Keiichi","family":"Okisawa","sequence":"additional","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6612-9339","authenticated-orcid":true,"given":"Hideki","family":"Taguchi","sequence":"additional","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]},{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9219-0726","authenticated-orcid":true,"given":"Takafumi","family":"Ueno","sequence":"additional","affiliation":[{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku , ,","place":["Yokohama, Japan, 226-8501"]},{"name":"Institute of Science Tokyo , , Nagatsuta-cho 4259, Midori-ku, ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,9,18]]},"reference":[{"issue":"24","key":"2026101210522873300_cit1","doi-asserted-by":"publisher","first-page":"17397","DOI":"10.1021\/acs.chemrev.2c00220","article-title":"Hierarchical Materials from High Information Content Macromolecular Building Blocks: Construction, Dynamic Interventions, and Prediction","volume":"122","author":"Shao","year":"2022","journal-title":"Chem. 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Protistol."},{"issue":"1","key":"2026101210522873300_cit25","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1128\/jb.168.1.40-48.1986","article-title":"Organization and Expression of Genetic Determinants for Synthesis and Assembly of Type 51 R Bodies","volume":"168","author":"Kanabrocki","year":"1986","journal-title":"J. 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