杨鹏难句41句的问题？

But achieving necessary matches in physical properties across interfaces between living and nonliving matter requires knowledge of which molecules control the bonding of cells to each other—an area that we have not yet explored thoroughly.

杨鹏难句的第41句中，主语matches为复数，而谓语requires用单数，这不对，可能是输入错误吧.他把of which 解释为which在此不是定语从句的引导词，而是一个形容词，意思是哪个，我认为不正确，请高手指导。