Optimization and variability can coexist

Many biological systems perform close to their physical limits, but promoting this optimality to a general principle seems to require implausibly fine tuning of parameters. Using examples from a wide range of systems, we show that this intuition is wrong. Near an optimum, functional performance depends on parameters in a "sloppy'' way, with some combinations of parameters being only weakly constrained. Absent any other constraints, this predicts that we should observe widely varying parameters, and we make this precise: the entropy in parameter space can be extensive even if performance on average is very close to optimal. This removes a major objection to optimization as a general principle, and rationalizes the observed variability.