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Chitwood, D. H.

Publications and source records attributed to Chitwood, D. H..

At least 19 recordsLinked to original sources

Disentangling blade and vasculature shape in grapevine leaves

The leaf blade and vasculature develop together within a shared morphological space. Despite shared molecular patterning pathways, it is unknown if developmental and evolutionary variation affect these tissues separately or together in a coordinated way. Grapevine leaves have a morphometric history and abundant data measuring the shape of the blade and vasculature together. Using a combination of topological data analysis and deep learning, we perform reciprocal semantic segmentation of leaf blade and vasculature. Each tissue contains sufficient information to predict the other. We hypothesize that this is due to a one-to-one relationship between blade and vein. Using thin plate splines to swap and warp different combinations of blade and vein shapes, we show that a set of leaves with a many-to-one relationship of blade and vein are distinguishable from true leaves. We also swap blade and vein across the developmental series and between species and show that only reversing the developmental series disrupts the relationship between blade and vasculature. We end by discussing the evolutionary and developmental implications that there is a unique, one-to-one mapping between blade and vein that allows each to be predicted from the other. Author summaryLeaves are made of two closely connected parts: the flat blade that captures light and the network of veins that transports water, nutrients, and developmental signals. Although these tissues grow together and share common molecular patterning pathways, it has remained unclear whether a particular blade shape is uniquely linked to a specific vein pattern. In this study, we use grapevine leaves as a model system and combine mathematical shape analysis with deep learning to examine this relationship. We show that the shape of the blade alone can accurately predict the vein network, and that the vein network can likewise predict the blade. This finding suggests a near one-to-one relationship between these two tissues. To test this idea, we created artificial leaves in which blade and vein shapes were deliberately mismatched. Although these synthetic leaves appeared realistic at a global level, a neural network was able to distinguish them from real leaves based on subtle differences. We further show that this tight coupling is maintained by the developmental sequence of leaf growth rather than by species identity, revealing a conserved constraint linking leaf form and internal structure.

plant biology↗

Procrustean pseudo-landmark methods in Python to measure massive quantities of leaf shape data

PremiseWhen examining leaf shapes that are different from one another, it can be difficult to compare both the overall leaf shape and points along the leaf margin in biologically and statistically meaningful ways. MethodTo address this problem, we present a simple and user-friendly leaf shape analysis in Jupyter Notebook and Python that uses pseudo-landmarks and Generalized Procrustes Analysis to measure and compare the shape of any leaf. To demonstrate our analysis, we created a repository of real leaves gathered from eight experimental datasets. ResultsUsing our leaf repository, we explain how we can use pseudo-landmarks to compare all leaf shapes both within and between species using dimension reduction techniques like Principal Component Analysis and can predict leaf shapes using pseudo-landmarks through Linear Discriminant Analysis. Our leaf shape analysis also maps differences in shape as leaves grew around a rosette, showing the transition of shape across development (phyllotaxy). Finally, we showed how we can investigate the relationship between leaf shape variation and genetic diversity by combining shape with genetic data. DiscussionThrough the use of Generalized Procrustes Analysis and pseudo-landmarks, our leaf shape analysis presents a powerful tool for examining the shape of any leaf across multiple biological, ecological, evolutionary, and developmental scales.

plant biology↗

Differences in apple fruit shape are independent of fruit size

PremiseFruit quality is crucial in breeding new apple accessions. Before tasting, consumers assess freshness and flavour based on the physical appearance of fruit. Understanding how fruit quality traits such as shape and size vary across diverse apples provides a foundation for future breeding efforts. MethodsWe analyzed images of 5724 apples representing 743 different trees and 534 unique accessions from Canadas Apple Biodiversity Collection to quantify variation in fruit shape and size. To achieve this, we used a pseudo-landmarking approach paired with traditional linear measurements including length, width, area, solidity, and aspect ratio. We also incorporated previously collected fruit weight measurements from the same trees. ResultsUsing a comprehensive measure of shape, we determined that the primary source of variation in apple fruit shape was the width to length (aspect) ratio of the fruit. This variation was not significantly correlated with differences in fruit size from its area and harvest weight. ConclusionsOur findings indicate that two critical aspects of morphological variation in apple--fruit shape and size--are independent, suggesting it is possible to select for a diverse range of fruit shapes while maintaining a consistent and marketable size.

plant biology↗

Interdisciplinarity through internationality: results from a US-Mexico graduate course bridging computational and plant science

Interdisciplinarity is used to integrate and synthesize new research directions between scientific domains, but it is not the only means by which to generate novelty by bringing diverse perspectives together. Internationality draws upon cultural and linguistic diversity that can potentially impact interdisciplinarity as well. We created an interdisciplinary class originally intended to bridge computational and plant science that eventually became international in scope, including students from the US and Mexico. We administered a survey over four years designed to evaluate student expertise. The first year of the survey included only US students and demonstrated that biology and computational student groups have distinct expertise but can learn the skills of the other group over the course of a semester. Modeling of survey responses shows that biological and computational science expertise is equally distributed between US and Mexico student groups, but that nonetheless these groups can be predicted based on survey responses due to sub-specialization within each domain. Unlike interdisciplinarity, differences arising from internationality are mostly static and do not change with educational intervention and include unique skills such as working across languages. We end by discussing a distinct form of interdisciplinarity that arises through internationality and the implications of globalizing research and education efforts.

scientific communication and education↗

Terroir and rootstock effects on leaf shape in California Central Valley vineyards

SummaryO_LIEmbedded 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. C_LIO_LIWe 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. C_LIO_LIUsing geometric morphometric approaches, we visualize a morphospace in which, in addition to clear separation of Cabernet Sauvignon and Chardonnay scion leaf shapes, an orthogonal source of shape variation affects both varieties. Comparing the Procrustes distances to within and between group means, the additional source of variance is found to arise from location and rootstock effects. C_LIO_LIWe describe and visualize a specific shape feature, the angle of the proximal lobe to the midvein that defines the closure of the petiolar sinus, that is attributable to location and rootstock effects and orthogonal to and separate from genetic, developmental, or allometric effects attributable to leaf size. C_LI Societal Impact Statement (EN)The 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 alter 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.

plant biology↗

A high resolution model of the grapevine leaf morphospace predicts synthetic leaves

O_LIGrapevine leaves are a model morphometric system. Sampling over ten thousand leaves using dozens of landmarks, the genetic, developmental, and environmental basis of leaf shape has been studied and a morphospace for the genus Vitis predicted. Yet, these representations of leaf shape fail to capture the exquisite features of leaves at high resolution. C_LIO_LIWe measure the shapes of 139 grapevine leaves using 1672 pseudo-landmarks derived from 90 homologous landmarks with Procrustean approaches. From hand traces of the vasculature and blade, we have derived a method to automatically detect landmarks and place pseudo-landmarks that results in a high-resolution representation of grapevine leaf shape. Using polynomial models, we create continuous representations of leaf development in 10 Vitis spp. C_LIO_LIWe visualize a high-resolution morphospace in which genetic and developmental sources of leaf shape variance are orthogonal to each other. Using classifiers, V. vinifera, Vitis spp., rootstock and dissected leaf varieties as well as developmental stages are accurately predicted. Theoretical eigenleaf representations sampled from across the morphospace that we call synthetic leaves can be classified using models. C_LIO_LIBy predicting a high-resolution morphospace and delimiting the boundaries of leaf shapes that can plausibly be produced within the genus Vitis, we can sample synthetic leaves with realistic qualities. From an ampelographic perspective, larger numbers of leaves sampled at lower resolution can be projected onto this high-resolution space; or, synthetic leaves can be used to increase the robustness and accuracy of machine learning classifiers. C_LI Societal Impact StatementGrapevine leaves are emblematic of the strong visual associations people make with plants. At a glance, leaf shape is immediately recognizable, and it is because of this reason it is used to distinguish grape varieties. In an era of computationally-enabled, machine learning-derived representations of reality, we can revisit how we view and use the shapes and forms that plants display to understand our relationship with them. Using computational approaches combined with time-honored methods, we can predict theoretical leaves that are possible to understand the genetics, development, and environmental responses of plants in new ways.

plant biology↗

Herbarium specimens reveal links between Capsella bursa-pastoris leaf shape and climate

O_LIStudies into the evolution and development of leaf shape have connected variation in plant form, function, and fitness. For species with consistent leaf margin features, patterns in leaf architecture are related to both biotic and abiotic factors. However, for species with inconsistent leaf margin features, quantifying leaf shape variation and the effects of environmental factors on leaf shape has proven challenging. C_LIO_LITo investigate leaf shape variation in species with inconsistent shapes, we analyzed approxi-mately 500 digitized Capsella bursa-pastoris specimens collected throughout the continental U.S. over a 100-year period with geometric morphometric modeling and deterministic techniques. We generated a morphospace of C. bursa-pastoris leaf shapes and modeled leaf shape as a function of environment and time. C_LIO_LIOur results suggest C. bursa-pastoris leaf shape variation is strongly associated with temperature over the C. bursa-pastoris growing season, with lobing decreasing as temperature increases. While we expected to see changes in variation over time, our results show that level of leaf shape variation is consistent over the 100-year period. C_LIO_LIOur findings showed that species with inconsistent leaf shape variation can be quantified using geometric morphometric modeling techniques and that temperature is the main environmental factor influencing leaf shape variation. C_LI

plant biology↗

Morphometrics and phylogenomics of coca (Erythroxylum spp.) illuminate its reticulate evolution, with implications for taxonomy

South American coca (Erythroxylum coca and E. novogranatense) has been a key-stone crop for many Andean and Amazonian communities for at least 8,000 years. However, over the last half century, global demand for cocaine has placed this plant in the centre of armed conflict, deforestation, and explosive growth of illegal economies. While national and international agencies progress from a war on drugs policy model towards locally appropriate, data-informed strategies to tackle coca plantations, monitoring their expansion and composition remains essential. The principal means to identify coca plants is leaf morphology, yet the extent to which it is reflected in taxonomy is uncertain. Here, we analyse the consistency of the current naming system of coca and its four closest wild relatives (the coca clade), using morphometrics, phylogenomics, and population genomics. We include the name-bearing type specimens of cocas closest wild relatives E. gracilipes and E. cataractarum. Morphometrics of 342 digitized herbarium specimens show that leaf shape and size fail to reliably discriminate between species and varieties. However, the rounder and more obovate leaves of certain coca varieties could be associated with domestication syndrome of this crop. Our phylogenomic data indicate gene flow involving monophyletic clades of E. gracilipes and the E. coca clade. These results further clarify the evolution of coca and support a taxonomic framework wherein E. gracilipes is retained as a single species. Our findings have implications for the development of cost-effective genotyping methods to effectively discriminate varieties of cultural significance from high-yielding cultivars fuelling the lucrative cocaine market.

plant biology↗

QTL analysis of floral morphology polymorphisms between Gilia yorkii and G. capitata

Speciation is a complex process typically accompanied by significant genetic and morphological differences between sister populations. In plants, this can result in divergent floral morphologies causing pollinator differences and reproductive isolation between populations. Here, we explore floral trait differences between two recently diverged species, Gilia yorkii and G. capitata. The distributions of floral traits in parental, F1, and F2 populations are compared, and groups of correlated traits are identified. We describe the genetic architecture of floral traits through a quantitative trait locus (QTL) analysis using an F2 population of 187 individuals. While all identified QTLs were of moderate (10-25%) effect, interestingly, many QTL intervals colocalized on Chromosomes 6 and 9, suggesting that sets of traits may share a common genetic basis. Our results provide a framework for future identification of genes involved in the evolution of floral morphology.

genetics↗

The shape and volume of air, kernels, and cracks, in a nutshell

Walnuts are the second most produced and consumed tree nut, with over 2.6 million metric tons produced in the 2022-23 harvest cycle alone. The United States is the second largest producer, accounting for 25% of the total global supply. Nonetheless, producers face an ever-growing demand in a more uncertain climate landscape, which requires effective and efficient walnut selection and breeding of new cultivars with increased kernel content and easy-to-open shells. Past and current efforts select for these traits using hand-held calipers and eye-based evaluations. Yet there is plenty of morphology that meets the eye but goes unmeasured, such as the volume of inner air or the convexity of the kernel. Here, we study the shape of walnut fruits based on X-ray CT (Computed Tomography) 3D reconstructions. We compute 49 different morphological phenotypes for 1264 individuals comprising 149 accessions. These phenotypes are complemented by traits of breeding interest such as ease of kernel removal and kernel weight. Through allometric relationships --relative growth of one tissue to another--, we identify possible biophysical constraints at play during development. We explore multiple correlations between all morphological and commercial traits, and identify which morphological traits can explain the most variability of commercial traits. We show that using only volume and thickness-based traits, especially inner air content, we can successfully encode several of the commercial traits. Core IdeasO_LIX-ray Computed Tomography (CT) imaging is used to compute a broad array of morpho-logical phenotypes in walnuts. C_LIO_LIThese morphological traits suggest biophysical constraints at play during walnut development. C_LIO_LIRelative inner air, shell, and packing tissue volumes are significantly correlated to the rest of shape phenotypes. C_LIO_LIThese volumes produce the best prediction models for traits of commercial interest such as shell strength. C_LIO_LIInexpensive phenotyping platforms that focus solely on volume measurement would enable better walnut breeding. C_LI

plant biology↗

From buds to shoots: Insights into grapevine development from the Witch's Broom bud sport

Premise of StudyDevelopment is relatively understudied in woody vines, such as grapevine (Vitis vinifera). We used the Witchs Broom bud sport in grapevine to understand the developmental trajectories of the bud sports, as well as the potential genetic basis of the bud sport. MethodsWe phenotyped shoots, buds, and leaves of two independent cases of the Witchs Broom bud sport, in the Dakapo and Merlot varieties of grapevine, alongside wild-type counterparts of the same variety. We also performed Illumina and Oxford Nanopore sequencing on the two independent cases and two wild-type counterparts of the same variety. Key ResultsThe Dakapo and Merlot cases of Witchs Broom displayed severe developmental defects, with no fruit/clusters formed and dwarf vegetative features. However, the Dakapo and Merlot cases of Witchs Broom studied were also phenotypically different from one another, with distinct differences in bud and leaf development. We were able to identify unique genetic mutations in our two Witchs Broom cases that are strong candidates for the genetic basis of the bud sports. ConclusionsThe Witchs Broom bud sport in grapevine serves as a useful natural mutant in which to study grapevine development. The Witchs Broom bud sports in both varieties studied had dwarf phenotypes, but the two instances studied were also vastly different from one another. Future work on Witchs Broom bud sports in grapevine could provide more insight into development and the genetic pathways involved in grapevine.

plant biology↗

Intra-leaf modeling of Cannabis leaflet shape produces synthetic leaves that predict genetic and developmental identities

O_LIThe iconic, palmately compound leaves of Cannabis have attracted significant attention in the past. However, investigations into the genetic basis of leaf shape or its connections to phytochemical composition have yielded inconclusive results. This is partly due to prominent changes in leaflet number within a single plant during development, which has so far prevented the proper use of common morphometric techniques. C_LIO_LIHere we present a new method that overcomes the challenge of nonhomologous landmarks in palmate, pinnate and lobed leaves, using Cannabis as an example. We model corresponding pseudo-landmarks for each leaflet as angle-radius coordinates and model them as a function of leaflet to create continuous polynomial models, bypassing the problems associated with variable number of leaflets between leaves. C_LIO_LIWe analyze 341 leaves from 24 individuals from nine Cannabis accessions. Using 3,591 pseudo-landmarks in modeled leaves, we accurately predict accession identity, leaflet number, and relative node number. C_LIO_LIIntra-leaf modeling offers a rapid, cost-effective means of identifying Cannabis accessions, making it a valuable tool for future taxonomic studies, cultivar recognition, and possibly chemical content analysis and sex identification, in addition to permitting the morphometric analysis of leaves in any species with variable numbers of leaflets or lobes. C_LI

developmental biology↗

Topological data analysis reveals a core heteroblastic program embedded in leaves of grapevine and maracuya

Leaves have a single shape. However, embedded in that shape are a multitude of latent shapes arising from evolutionary, developmental, environmental, and other effects. These confounded effects manifest at distinct developmental time points and evolve at different tempos. Here, revisiting datasets comprised of thousands of leaves of vining grapevine (Vitaceae) and maracuya (Passifloraceae) species, we apply a technique from the mathematical field of topological data analysis to comparatively visualize the structure of heteroblastic effects on leaf shape in each group. Consistent with a morphologically closer relationship, members of the grapevine dataset possess a strong core heteroblasty program with little deviation between species. Remarkably, we found that most members of the maracuya family also share a core heteroblasty program despite dramatic species-to-species leaf shape differences. This conservation was not detected using traditional analyses. We also identify two morphotypes of maracuya that deviate from the core structure, suggesting the evolution of new heteroblastic properties in this phylogenetically related sub-group. Our findings illustrate how topological data analysis can be used to disentangle previously confounded developmental and evolutionary effects to visualize latent shapes and hidden relationships, even ones embedded in complex, high-dimensional datasets.

plant biology↗

Integrated PET and confocal imaging informs a functional timeline for the dynamic process of vascular reconnection during grafting.

Grafting is a widely used agricultural technique that involves the physical joining of separate plant parts so they form a unified vascular system, enabling beneficial traits from independent genotypes to be captured in a single plant. This simple, yet powerful tool has been used for thousands of years to improve abiotic and biotic stress tolerance, enhance yield, and alter plant architecture in diverse crop systems. Despite the global importance and ancient history of grafting, our understanding of the fundamental biological processes that make this technique successful remains limited, making it difficult to efficiently expand on new genotypic graft combinations. One of the key determinants of successful grafting is the formation of the graft junction, an anatomically unique region where xylem and phloem strands connect between newly joined plant parts to form a unified vascular system. Here, we use an integrated imaging approach to establish a spatiotemporal framework for graft junction formation in the model crop Solanum lycopersicum (tomato), a plant that is commonly grafted worldwide to boost yield and improve abiotic and biotic stress resistance. By combining Positron Emission Tomography (PET), a technique that enables the spatio-temporal tracking of radiolabeled molecules, with high-resolution laser scanning confocal microscopy (LSCM), we are able to merge detailed, anatomical differentiation of the graft junction with a quantitative timeline for when xylem and phloem connections are functionally re-established. In this timeline, we identify a 72-hour window when anatomically connected xylem and phloem strands regain functional capacity, with phloem restoration typically preceding xylem restoration by about 24-hours. Furthermore, we identify heterogeneity in this developmental and physiological timeline that corresponds with microvariability in the physical contact between newly joined rootstock-scion tissues. Our integration of PET and confocal imaging technologies provides a spatio-temporal timeline that will enable future investigations into cellular and tissue patterning events that underlie successful versus failed vascular restoration across the graft junction.

plant biology↗

Global disparities in plant science: a legacy of colonialism, patriarchy, and exclusion

The field of plant science has grown dramatically in the past two decades, but global disparities and systemic inequalities persist. Here, we analyzed ~300,000 papers published over the past two decades to quantify disparities across nations, genders, and taxonomy in the plant science literature. Our analyses reveal striking geographical biases--affluent nations dominate the publishing landscape and vast areas of the globe having virtually no footprint in the literature. Authors in Northern America are cited nearly twice as many times as authors based in Sub-Saharan Africa and Latin America, despite publishing in journals with similar impact factors. Gender imbalances are similarly stark and show remarkably little improvement over time. Some of the most affluent nations have extremely male biased publication records, despite supposed improvements in gender equality. In addition, we find that most studies focus on economically important crop and model species and a wealth of biodiversity is under-represented in the literature. Taken together, our analyses reveal a problematic system of publication, with persistent imbalances that poorly captures the global wealth of scientific knowledge and biological diversity. We conclude by highlighting disparities that can be addressed immediately and offer suggestions for long-term solutions to improve equity in the plant sciences. SIGNIFICANCE STATEMENTWe analyzed ~300,000 papers published over the past two decades to quantify global, gender, and taxonomic disparities in plant science. Our analyses reveal striking geographical biases that are correlated with national affluence. Gender imbalances were also evident, with far more papers led by authors with masculine names than authors with feminine names. Lastly, we identified substantial taxonomic sampling gaps. The vast majority of surveyed studies focused on major crop and model species and the remaining biodiversity accounted for only a fraction of publications. Taken together, our analyses represent an important addition to the growing conversation about diversifying and decolonizing science.

plant biology↗

The topological shape of gene expression across the evolution of flowering plants

Since they emerged ~125 million years ago, flowering plants have evolved to dominate the terrestrial landscape and survive in the most inhospitable environments on earth. At their core, these adaptations have been shaped by changes in numerous, interconnected pathways and genes that collectively give rise to emergent biological phenomena. Linking gene expression to morphological outcomes remains a grand challenge in biology, and new approaches are needed to begin to address this gap. Here, we implemented topological data analysis (TDA) to summarize the high dimensionality and noisiness of gene expression data using lens functions that delineate plant tissue and stress responses. Using this framework, we created a topological representation of the shape of gene expression across plant evolution, development, and environment for the phylogenetically diverse flowering plants. The TDA-based Mapper graphs form a well-defined gradient of tissues from leaves to seeds, or from healthy to stressed samples, depending on the lens function. This suggests there are distinct and conserved expression patterns across angiosperms that delineate different tissue types or responses to biotic and abiotic stresses. Genes that correlate with the tissue lens function are enriched in central processes such as photosynthetic, growth and development, housekeeping, or stress responses. Together, our results highlight the power of TDA for analyzing complex biological data and reveal a core expression backbone that defines plant form and function. Significance statementA grand challenge in biology is to link gene expression to phenotypes across evolution, development, and the environment, but efforts have been hindered by biological complexity and dataset heterogeneity. Here, we implemented topological data analysis across thousands of gene expression datasets in phylogenetically diverse flowering plants. We created a topological representation of gene expression across plants and observed well-defined gradients of tissues from leaves to seeds, or from healthy to environmentally stressed. Using this framework, we identified a core and deeply conserved expression backbone that defines plant form and function, with key patterns that delineate plant tissues, abiotic, and biotic stresses. Our results highlight the power of topological approaches for analyzing complex biological datasets.

plant biology↗

Grapevine leaf size influences vine canopy temperature

PremiseGrapevine leaves have diverse shapes and sizes. Their shape and size is known to be influenced by many factors including genetics, vine phytosanitary status, environment, leaf and vine age, and node position on the shoot. In order to determine the importance of grapevine leaf shape and size to canopy temperature, we examined the relationship in five seedling populations grown in a vineyard in California, USA. MethodsAll of the populations had one parent with compound leaves of the Vitis piasezkii type and each population had a different second parent with non-compound leaves. In previous work, we measured leaf shape and size using 21 homologous landmarks. Here, we paired these morphology data with measurements taken using an infrared thermometer to measure the temperature of the canopy. By recording time of sampling and canopy temperature, we were able to determine which vines were cooler or hotter than expected, using a linear model. ResultsWe established a relationship between leaf size and canopy temperature: vines with larger leaves were cooler than expected. In contrast, leaf shape was not strongly correlated with variation in temperature. ConclusionsUltimately, these findings indicate that vines with larger leaves may contribute to the reduction of overall vine canopy temperature, but further work is needed to determine if this is due to variation in leaf size, differences in the openness of the canopy, or other related traits.

plant biology↗

X-ray imaging of 30 year old wine grape wood reveals cumulative impacts of rootstocks on scion secondary growth and harvest index

O_LIAnnual rings from 30 year old vines in a California rootstock trial were measured to determine the effects of 15 different rootstocks on Chardonnay and Cabernet Sauvignon scions. Viticultural traits measuring vegetative growth, yield, berry quality, and nutrient uptake were collected at the beginning and end of the lifetime of the vineyard. C_LIO_LIX-ray Computed Tomography (CT) was used to measure ring widths in 103 vines. Ring width was modeled as a function of ring number using a negative exponential model. Early and late wood ring widths, cambium width, and scion trunk radius were correlated with 27 traits. C_LIO_LIModeling of annual ring width shows that scions alter the width of the first rings but that rootstocks alter the decay thereafter, consistently shortening ring width throughout the lifetime of the vine. The ratio of yield to vegetative growth, juice pH, photosynthetic assimilation and transpiration rates, and stomatal conductance are correlated with scion trunk radius. C_LIO_LIRootstocks modulate secondary growth over years, altering hydraulic conductance, physiology, and agronomic traits. Rootstocks act in similar but distinct ways from climate to modulate ring width, which borrowing techniques from dendrochronology, can be used to monitor both genetic and environmental effects in woody perennial crop species. C_LI

plant biology↗