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Societal Impact StatementThe innumerable effects of terroir—including climate, soil, microbial environment, biotic interactions, and cultivation practice—collectively alter plant performance and production. A more direct agricultural intervention is grafting, in which genetically distinct shoot and root genotypes are surgically combined to create a chimera that alters shoot performance at a distance. Selection of location and rootstock are intentional decisions in viticulture to positively alter production outcomes. Here, we show that terroir and rootstock alter the shapes of grapevine leaves in commercial vineyards throughout the California Central Valley, documenting the profound effects of these agricultural interventions that alter plant morphology. SummaryEmbedded in a single leaf shape are the latent signatures of genetic, developmental, and environmental effects. In viticulture, choice of location and rootstock are important decisions that affect the performance and production of the shoot. We hypothesize that these effects influence plant morphology, as reflected in leaf shape.We sample 1879 leaves arising from scion and rootstock combinations from commercial vineyards in the Central Valley of California. Our design tests 20 pairwise contrasts between Cabernet Sauvignon and Chardonnay scions from San Joaquin, Merced, and Madera counties from vines grafted to Teleki 5C, 1103 Paulsen, and Freedom rootstocks.We quantify clear differences between Cabernet Sauvignon and Chardonnay leaves. However, we also detect a separate, statistically independent source of shape variance that affects both Cabernet Sauvignon and Chardonnay leaves similarly. We find that this other shape difference is associated with differences in rootstock and location.The shape difference that arises from rootstock and location affects the basal part of the leaf near the petiole, known as the petiolar sinus, and affects its closure. This shape effect is independent from previously described shape differences that arise from genetic, developmental, or size effects. 

Grapevine leaves have diverse shapes and sizes which are influenced by many factors including genetics, vine phytosanitary status, environment, leaf and vine age, and node position on the shoot. To determine the relationship between grapevine leaf shape or size and leaf canopy temperature, we examined five seedling populations grown in a vineyard in California, USA. The populations had one parent with compound leaves of the Vitis piasezkii type and a different second parent with non-compound leaves. In previous work, we had measured the shape and size of the leaves collected from these populations using 21 homologous landmarks. Here, we paired these morphological data with canopy temperature measurements made using a handheld infrared thermometer. After recording time of sampling and canopy temperature, we used a linear model between time of sampling and canopy temperature to estimate temperature residuals. Based on these residuals, we determined if the canopy temperature of each vine was cooler or warmer than expected, based on the time of sampling. We established a relationship between leaf size and canopy temperature: vines with larger leaves were cooler than expected. By contrast, leaf shape was not strongly correlated with variation in canopy temperature. Ultimately, these findings indicate that vines with larger leaves may contribute to the reduction of overall canopy temperature; however, further work is needed to determine whether this is due to variation in leaf size, differences in the openness of the canopy or other related traits. 

Indirect defenses are plant phenotypes that reduce damage by attracting natural enemies of plant pests and pathogens to leaves. Despite their economic and ecological importance, few studies have investigated the genetic underpinnings of indirect defense phenotypes. Here, we present a genome-wide association study of five phenotypes previously determined to increase populations of beneficial (fungivorous and predacious) mites on grape leaves (genus Vitis): leaf bristles, leaf hairs, and the size, density, and depth of leaf domatia. Using a common garden genetic panel of 399 V. vinifera cultivars, we tested for genetic associations of these phenotypes using previously obtained genotyping data from the Vitis9kSNP array. We found one single nucleotide polymorphism (SNP) significantly associated with domatia density. This SNP (Chr5:1160194) is near two genes of interest: Importin Alpha Isoform 1 (VIT_205s0077g01440), involved in downy mildew resistance, and GATA Transcription Factor 8 (VIT_205s0077g01450), involved in leaf shape development. Our findings are among the first to examine the genomic regions associated with ecologically important plant traits that facilitate interactions with beneficial mites, and suggest promising candidate genes for breeding and genetic editing to increase naturally occurring predator-based defenses in grapevines. 

PremiseLeaf morphology is dynamic, continuously deforming during leaf expansion and among leaves within a shoot. Here, we measured the leaf morphology of more than 200 grapevines (Vitisspp.) over four years and modeled changes in leaf shape along the shoot to determine whether a composite leaf shape comprising all the leaves from a single shoot can better capture the variation and predict species identity compared with individual leaves. MethodsUsing homologous universal landmarks found in grapevine leaves, we modeled various morphological features as polynomial functions of leaf nodes. The resulting functions were used to reconstruct modeled leaf shapes across the shoots, generating composite leaves that comprehensively capture the spectrum of leaf morphologies present. ResultsWe found that composite leaves are better predictors of species identity than individual leaves from the same plant. We were able to use composite leaves to predict the species identity of previously unassigned grapevines, which were verified with genotyping. DiscussionObservations of individual leaf shape fail to capture the true diversity between species. Composite leaf shape—an assemblage of modeled leaf snapshots across the shoot—is a better representation of the dynamic and essential shapes of leaves, in addition to serving as a better predictor of species identity than individual leaves.