where $\sigma_G^2$ was the genotypic variance calculated based on the above models with a random effect for $G_i$ and $\bar{\upsilon}_\delta^BLUE$ was the mean variance of the difference of two genotypic means (Holland et al., 2003; Piepho and Möhring, 2007).
To avoid the effect of the varying number of replicates, the $H^2$ of photosynthesis-related parameters was estimated for each developmental stage based on the following equation: