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. 2022 Jan 3;8:798353. doi: 10.3389/fmolb.2021.798353

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Copyright © 2022 Padavannil, Ayala-Hernandez, Castellanos-Silva and Letts.

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

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Accessory subunits of the CI heel. The structure of CI (PDB: 6ZKC used throughout) in cartoon with the surface of NDUFA13 (light yellow) (A), NDUFA3 (pine green) (B) and NDUFA1 (firebrick red) (C) shown, the core subunits colored as in Figure 1A and the accessory subunits grey. (D) Looking up from the CI heel showing NDUFA13, NDUFA3, and NDUFA1 of H. sapiens CI structure (PDB: 5XTD) as cartoons. NDUFA8 (lilac purple) is shown as a transparent surface for clarity. (E) Accessory subunit NDUFA13 of H. sapiens CI structure (PDB: 5XTD) is shown as cartoon. The residues mentioned in the text are shown in stick representation. (F) NDUFA13 compensates for the lost transmembrane helix TMH9^ND1 (cobalt blue) of T. thermophilus. T. thermophilus ND1 (Nqo8) (PDB: 4HEA used throughout) is superposed on ND1 of mammalian (O. aries) CI structure shown as transparent surface. TMH5^ND1(Nqo8) and TMH9 ^ND1(Nqo8) are shown as cartoons. NDUFA13 surface and cartoon are shown. (G) Schematic representation of the “Flag post model.” NDUFA13 of healthy CI is furled and incapable of binding to STAT3. NDUFA13 alone is raised exposing the STAT3 interacting site to the IMS. (H,I) Assembly vs. lipid binding roles of NDUFA3. (H) Mammalian CI shown as transparent surface with NDUFA3, NDUFA13, α1^ND1 (cobalt blue), and α3^ND3 (sea green) shown as cartoons. (I) Y. lipolytica CI (PDB: 6YJ4) shown as transparent surface with NDUFA3 (NI9M), NDUFA13 (NB6M), NUXM (light cyan), α1^ND1 (NU1M), and α3^ND3 (NU3M) (sea green) shown as cartoons. (J,K) NDUFA1 compensates for the shorter TMH2-3^ND1 loop. (J) Mammalian CI structure shown as transparent surface with NDUFA1, TMH1^ND1, TMH7^ND1 and the TMH2-3^ND1 loop shown as cartoons. (K) T. thermophilus CI shown as transparent surface with TMH1^ND1(Nqo8), TMH7^ND1(Nqo8) and the TMH2-3^ND1(Nqo8) loop shown as cartoon. (L) NDUFA1 connects the P_P-a and P_P-b subassemblies. Mammalian CI a transparent surface with NDUFA1 and the ND6 TMH4-5^ND6 (cyan-blue) loop shown as cartoon. Residues mentioned in the text are shown as sticks. Key H40-S46 sequence shown in purple. Subunits are colored as in Figure 1 in all panels unless stated otherwise. NDUFA13: light yellow, NDUFA3: pine, NDUFA1: firebrick red, NDUFA8: lilac purple, ND1: cobalt blue, ND3: sea green, ND6: cyan-blue, NUXM (light cyan).