Your paper is very interesting. In 2015, by computing the frameshift substitution scores, we have found that the natural standard genetic code was optimized for frameshift tolerance. Here you analyzed the frameshift tolerance of the genetic codes in a different way. The result is basically consistent with ours, and seems better in some aspects.
Recently we have further analyzed the frameshift tolerance of the genomes of some model species, and revealed the molecular mechanism of frameshift repair.
If you are interested, you may read our papers. Welcome any comments and suggestions!
Xiaolong Wang, et al. (2015) The shiftability of protein coding genes: the genetic code was optimized for frameshift tolerating. PeerJ PrePrints 3: e806v1 http://dx.doi.org/10.7287/peerj.preprints.806v1
Xiaolong Wang, Quanjiang Dong, Gang Chen, Jianye Zhang, Yongqiang Liu, Jinqiao Zhao, Haibo Peng, Yalei Wang, Yujia Cai, Xuxiang Wang, Chao Yang, Michael Lynch, The universal genetic code, protein coding genes and genomes of all species were optimized for frameshift tolerance, bioRxiv 067736; doi: https://doi.org/10.1101/067736
Xiaolong Wang, Haibo Peng, Chunyan Li, Xuxiang Wang, Yalei Wang, Gang Chen, Jianye Zhang, Michael Lynch, Premature termination codons signal targeted gene repair by nonsense mRNA-mediated gene editing in E. coli, bioRxiv 069971; doi: https://doi.org/10.1101/069971
College of Life Sciences,
Ocean University of China