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. 2017 Aug 25;7:9558. doi: 10.1038/s41598-017-09749-2

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© The Author(s) 2017

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Schematic of inference and validation of the α-helix rule and hydropathy contradiction rule. For the α-helix rule, we made inferences based on positive variants of four enzymes obtained by random mutagenesis screening. The rule was inferred by saturating mutagenesis at these sites in ChMOX (Val455) and AtADC (Leu435). On the other hand, for the hydropathy contradiction rule, we made inferences based on sequence analysis of MeHNL. The rule was validated by identifying five candidate hotspots in SuPDH. Finally, we evaluated the effectiveness of the rules by applying the method to MpLUC and hGH.