Which of the following was used as evidence by Dingemanse and colleagues (2002) to conclude that exploratory behavior in great tits was a heritable trait?

A) Selection on exploratory behavior resulted in strong population changes over four generations
B) Individuals differed greatly in their exploratory behavior
C) Inbreeding did not affect exploratory behavior
D) Individuals with high exploratory behavior had the highest fitness
E) The exploratory behavior of an individual was highly repeatable