Symmetries of the defocusing nonlinear Schrödinger equation are expressed in action-angle coordinates and characterized in terms of the periodic and Dirichlet spectrum of the associated Zakharov-Shabat system. Application: proof of the conjecture that the periodic spectrum $\cdots \<{\lambda }_k^{-}\le {\lambda }_k^{+}\<{\lambda }_{k+1}^{-}\le \cdots$ of a Zakharov-Shabat operator is symmetric,i.e. ${\lambda }_k^{\pm }=-{\lambda }_{-k}^{\mp }$ for all $k$, if and only if the sequence ${\left({\gamma }_k\right)}_{k\in \mathbb{Z}}$ of gap lengths, ${\gamma }_k:={\lambda }_k^{+}-{\lambda }_k^{-}$, is symmetric with respect to $k=0$.