Genetic architecture of leaf morphology revealed by integrated trait module in Catalpa bungei

Leaves are crucial for maintaining plant growth and development via photosynthesis,and their function is simultaneously regulated by a suite of phenotypic traits.Although much is known about the genetic architecture of individual leaf traits,unraveling the genetic basis of complex leafmorphology remains a challenge.Based on the functional correlation and coordination ofmulti-traits,we divided 15 leaf morphological traits into three modules,comprising size(area,length,width,and perimeter),shape(leaf lobes,aspect ratio,circularity,rectangularity,and the relevant ratios),and color(red,green,and blue)for an ornamental tree species,Catalpa bungei.A total of 189 significant single-nucleotide polymorphisms were identified in the leaves of C.bungei:35,82,and 76 in the size,shape,and color modules,respectively.Four quantitative trait loci were common between the size and shape modules,which were closely related according to phenotype correlation,genetic mapping,and mRNA analysis.The color module was independent of them.Synergistic changes in the aspect ratio,leaf lobe,and circularity suggest that these traits could be the core indicators of the leaf shape module.The LAS and SRK genes,associated with leaf lobe and circularity,were found to function in plant defense mechanisms and the growth of leaves.The associations between the SRK and CRK2 genes and the leaf lobe and circularity traits were further verified by RT-qPCR.Our findings demonstrate the importance of integrating multi-trait modules to characterize leaf morphology and facilitate a holistic understanding of the genetic architecture of intraspecific leaf morphology diversity.