What Is the Major Problem Associated with the Application of Optimality

What is the major problem associated with the application of optimality theory to the study of animal behavior?

A) It is virtually impossible to find a common unit in which to translate all costs and benefits.
B) It is easy to measure the benefits of a behavior but very difficult to measure its costs.
C) It is not possible to determine the number of offspring that might have existed if the parent had behaved in a different fashion.
D) all of the above.

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