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The insecticidal crystal protein (Cry) genes of  Bacillus thuringiensis  are a key gene resource for generating transgenic crops with pest resistance. However, many  cry  genes cannot be expressed or form crystals in mother cells. Here, we report a novel Cry protein gene,  cry65Aa1 , which exists in an operon that contains a downstream gene encoding a hypothetical protein ORF2. We demonstrated that ORF2 is required for Cry65Aa1 expression and crystallization by function as a C-terminal crystallization domain. The  orf2  sequence is also required for Cry65Aa expression, because  orf2  transcripts have a stabilizing effect on  cry65Aa1  transcripts. Furthermore, we found that the crystallization of Cry65Aa1 required the Cry65Aa1 C-terminus in addition to ORF2 or a typical Cry protein C-terminal region. Finally, we showed that Cry65Aa1 has a selective cytotoxic effect on MDA-MB231 cancer cells. This report is the first description of a 130-kDa mass range Cry protein requiring two C-termini for crystallization. Our findings reveal a novel evolutionary strategy of Cry proteins and provide an explanation for the existence of Cry protein genes that cannot form crystals in  B. thuringiensis . This study also provides a potential framework for isolating novel  cry  genes from “no crystal”  B. thuringiensis  strains.

The expression and crystallization of Cry65Aa require two C-termini, revealing a novel evolutionary strategy of Bacillus thuringiensis Cry proteins