Optimal isn't good enough.

The notion that biological systems come to embody optimal solutions seems consistent with the competitive drive of evolution. It has been used to interpret many examples of sensorimotor behavior. It is attractive from the viewpoint of control engineers because it solves the redundancy problem by identifying the one optimal motor strategy out of many similarly acceptable possibilities. This perspective examines whether there is sufficient basis to apply the formal engineering tools of optimal control to a reductionist understanding of biological systems. For an experimental biologist, this translates into whether the theory of optimal control generates nontrivial and testable hypotheses that accurately predict novel phenomena, ideally at deeper levels of structure than the observable behavior. The methodology of optimal control is applicable when there is (i) a single, known cost function to be optimized, (ii) an invertible model of the plant, and (iii) simple noise interfering with optimal performance. None of these is likely to be true for biological organisms. Furthermore, their motivation is usually good-enough rather than globally optimal behavior. Even then, the performance of a biological organism is often much farther from optimal than the physical limits of its hardware because the brain is continuously testing the acceptable limits of performance as well as just performing the task. This perspective considers an alternative strategy called "good-enough" control, in which the organism uses trial-and-error learning to acquire a repertoire of sensorimotor behaviors that are known to be useful, but not necessarily optimal. This leads to a diversity of solutions that tends to confer robustness on the individual organism and its evolution. It is also more consistent with the capabilities of higher sensorimotor structures, such as cerebral cortex, which seems to be designed to classify and recall complex sets of information, thereby allowing the organism to learn from experience, rather than to compute new strategies online. Optimal control has been a useful metaphor for understanding some superficial aspects of motor psychophysics. Reductionists who want to understand the underlying neural mechanisms need to move on.