Morphometrics Reveals Complex And Heritable Apple Leaf Shapes

Apple (Malus spp.) is a widely grown and valuable fruit crop. Leaf shape and size are important for flowering in apple and may also be early indicators for other agriculturally valuable traits. We examined 9,000 leaves from 869 unique apple accessions using linear measurements and comprehensive morphometric techniques. We identified allometric variation in the length-to-width aspect ratio between accessions and species of apple. The allometric variation was due to variation in the width of the leaf blade, not length. Aspect ratio was highly correlated with the primary axis of morphometric variation (PC1) quantified using elliptical Fourier descriptors (EFDs) and persistent homology (PH). While the primary source of variation was aspect ratio, subsequent PCs corresponded to complex shape variation not captured by linear measurements. After linking the morphometric information with over 122,000 genome-wide SNPs, we found high narrow-sense heritability values even at later PCs, indicating that comprehensive morphometrics can capture complex, heritable phenotypes. Thus, techniques such as EFDs and PH are capturing heritable biological variation that would be missed using linear measurements alone, and which could potentially be used to select for a hidden phenotype only detectable using comprehensive morphometrics.