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The mitophagy receptor Nix interacts with LC3/GABARAP proteins, targeting mitochondria into autophagosomes for degradation. Here we present evidence for phosphorylation-driven regulation of the Nix:LC3B interaction. Isothermal titration calorimetry and NMR indicate a ~100 fold enhanced affinity of the serine 34/35-phosphorylated Nix LC3-interacting region (LIR) to LC3B and formation of a very rigid complex compared to the non-phosphorylated sequence. Moreover, the crystal structure of LC3B in complex with the Nix LIR peptide containing glutamic acids as phosphomimetic residues and NMR experiments revealed that LIR phosphorylation stabilizes the Nix:LC3B complex via formation of two additional hydrogen bonds between phosphorylated serines of Nix LIR and Arg11, Lys49 and Lys51 in LC3B. Substitution of Lys51 to Ala in LC3B abrogates binding of a phosphomimetic Nix mutant. Functionally, serine 34/35 phosphorylation enhances autophagosome recruitment to mitochondria in HeLa cells. Together, this study provides cellular, biochemical and biophysical evidence that phosphorylation of the LIR domain of Nix enhances mitophagy receptor engagement.
This paper is intended to investigate the linguistic behaviors of the Korean as-parenthetical constructions with the aim of devoting to distinguishing universal properties of as-parentheticals. This paper shows three prominent behaviors in Korean as-parenthetical constructions. First, the Korean as-clause displays that the syntactic gap in as-clauses must be realized as CP, through the variations on case marker. Secondly, the Korean as-parentheticals tend to have two types of as-clauses; CP or VP as-clause types. In addition, they are sensitive to the syntactic restrictions which can be noticed in as-parenthetical constructions: the sisterhood restriction and the Island boundary. Thirdly, the Korean as-parenthetical constructions reveal that they would require some pragmatic information which is combined with semantic meaning, in the process of getting the interpretation of as-clauses.