“…Q d is suitable for analyzing metabolic networks because the networks are directed (Clune et al, 2013). For example, unlike Q ud , Q d can characterize input (i.e., nutrient) modules, core modules, and output (i.e., product) modules (Leicht & Newman, 2008) (e.g., a bow-tie structure, which is related to biological robustness (Friedlander, Mayo, Tlusty, & Alon, 2014;Ma & Zeng, 2003)) in metabolic networks by considering edge direction. Nonetheless, some previous studies (Parter et al, 2007;Takemoto & Borjigin, 2011;Takemoto, 2013;Wagner & Fell, 2001) focused on the undirected version of the Q-value because of some apparent problems with Q d ; for example, the reversibility/irreversibility of metabolic reactions may change with environmental conditions (Parter et al, 2007;Wagner & Fell, 2001), and Q d cannot distinguish between situations with and without directed flow (Arenas, Duch, Fernández, & Gómez, 2007) (e.g., Fig.…”