The proton and metal binding sites responsible for the pH-dependent green-red bioluminescence color tuning in firefly luciferases.

In the pH-sensitive luciferases and their mutants, although the calculation of the overall microenvironment charge, considering the four residues H310, E311, R337 and E354, showed a trend for blue-shifted emission in electrostatically more negative environments and red-shifted emission for more positive environments (Table 2 ), the mutant R337E, in which there is a substantial increase of the overall negative charge character of the environment, also resulted in red light.

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