Biology subjects

Chitwood, D. H.

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

At least 37 records · Page 2Linked to original sources

bioRxiv2022

The shape of aroma: measuring and modeling citrus oil gland distribution

O_LICitrus come in diverse sizes and shapes, and play a key role in world culture and economy. Citrus oil glands in particular contain essential oils which include plant secondary metabolites associated with flavor and aroma. Capturing and analyzing nuanced information behind the citrus fruit shape and its oil gland distribution provides a morphology-driven path to further our insight into phenotype-genotype interactions. C_LIO_LIWe investigated the shape of citrus fruit of 51 accessions based on 3D X-ray CT scan reconstructions. Accessions include all three ancestral citrus species, accessions from related genera, and several interspecific hybrids. We digitally separate and compare the size of fruit endocarp, mesocarp, exocarp, and oil gland tissue. Based on the centers of the oil glands, overall fruit shape is approximated with an ellipsoid. Possible oil gland distributions on this ellipsoid surface are explored using directional statistics. C_LIO_LIThere is a strong allometry along fruit tissues; that is, we observe a strong linear relationship between the volume of any pair of major tissues. This suggests that the relative growth of fruit tissues with respect to each other follows a power law. We also observe that on average, glands distance themselves from their nearest neighbor following a square root relationship, which suggests normal diffusion dynamics at play. C_LIO_LIThe observed allometry and square root models point to the existence of biophysical developmental constraints that govern novel relationships between fruit dimensions from both evolutionary and breeding perspectives. Understanding these biophysical interactions prompt an exciting research path on fruit development and breeding. C_LI Societal Impact StatementCitrus are intrinsically connected to human health and culture, including preventing human diseases like scurvy, and inspiring sacred rituals. Citrus fruits come in a stunning number of different sizes and shapes, ranging from small clementines to oversized pummelos, and fruits display a vast diversity of flavors and aromas. These qualities are key in both traditional and modern medicine and the production of cleaning and perfume products. By quantifying and modeling overall fruit shape and oil gland distribution, we can gain further insight into citrus development and the impacts of domestication and improvement on multiple characteristics of the fruit.

plant biology↗

bioRxiv2022

Increases in vein length compensate for leaf area lost to lobing in grapevine

PremiseThere is considerable variation in leaf lobing and leaf size, including among grapevines, some of the most well-studied leaves. We examined the relationship between leaf lobing and leaf size across grapevine populations which varied in extent of leaf lobing. MethodsWe used homologous landmarking techniques to measure 2,632 leaves across two years in 476 unique, genetically distinct grapevines from 5 biparental crosses which vary primarily in the extent of lobing. We determined to what extent leaf area could explain variation in lobing, vein length, and vein to blade ratio. ResultsAlthough lobing was the primary source of variation in shape across the leaves we measured, leaf area varied only slightly as a function of lobing. Rather, leaf area increases as a function of total major vein length, total branching vein length, and decreases as a function of vein to blade ratio. These relationships are stronger for more highly lobed leaves, with the residuals for each model differing as a function of distal lobing. ConclusionsFor a given leaf area, more highly lobed leaves have longer veins and higher vein to blade ratios, allowing them to maintain similar leaf areas despite increased lobing. These findings show how more highly lobed leaves may compensate for what would otherwise result in a reduced leaf area, allowing for increased photosynthetic capacity through similar leaf size.

plant biology↗

bioRxiv2022

The grapevine leaves that are (or were) not: constraints on leaf development and choosing to see what seemingly is not

Using conventional statistical approaches there exist powerful methods to classify shapes. Embedded in morphospaces is information that allows us to visualize theoretical leaves. These unmeasured leaves are never considered nor how the negative morphospace can inform us about the forces responsible for shaping leaf morphology. Here, we model leaf shape using an allometric indicator of leaf size, the ratio of vein to blade areas. The borders of the observable morphospace are restricted by constraints and define an orthogonal grid of developmental and evolutionary effects which can predict the shapes of possible grapevine leaves. Leaves in the genus Vitis are found to fully occupy morphospace available to them. From this morphospace we predict the developmental and evolutionary shapes of grapevine leaves that are not only possible, but exist, and argue that rather than explaining leaf shape in terms of discrete nodes or species, that a continuous model is more appropriate.

plant biology↗

bioRxiv2021

Measuring hidden phenotype: Quantifying the shape of barley seeds using the Euler Characteristic Transform

Shape plays a fundamental role in biology. Traditional phenotypic analysis methods measure some features but fail to measure the information embedded in shape comprehensively. To extract, compare, and analyze this information embedded in a robust and concise way, we turn to Topological Data Analysis (TDA), specifically the Euler Characteristic Transform. TDA measures shape comprehensively using mathematical representations based on algebraic topology features. To study its use, we compute both traditional and topological shape descriptors to quantify the morphology of 3121 barley seeds scanned with X-ray Computed Tomography (CT) technology at 127 micron resolution. The Euler Characteristic Transform measures shape by analyzing topological features of an object at thresholds across a number of directional axes. A Kruskal-Wallis analysis of the information encoded by the topological signature reveals that the Euler Characteristic Transform picks up successfully the shape of the crease and bottom of the seeds. Moreover, while traditional shape descriptors can cluster the seeds based on their accession, topological shape descriptors can cluster them further based on their panicle. We then successfully train a support vector machine (SVM) to classify 28 different accessions of barley based exclusively on the shape of their grains. We observe that combining both traditional and topological descriptors classifies barley seeds better than using just traditional descriptors alone. This improvement suggests that TDA is thus a powerful complement to traditional morphometrics to comprehensively describe a multitude of "hidden" shape nuances which are otherwise not detected.

plant biology↗

bioRxiv2021

Quantitative dissection of color patterning in the foliar ornamental Coleus reveals underlying features driving aesthetic value

O_LIColeus is a popular ornamental plant that exhibits a diverse array of foliar color patterns. New cultivars are currently hand selected by both amateur and experienced plant breeders. In this study, we reimagine coleus breeding using a quantitative color analysis framework. C_LIO_LIDespite impressive advances in high-throughput data collection and processing, complex color patterns remain challenging to extract from image datasets. Using a new phenotyping approach called "ColourQuant," we extract and analyze pigmentation patterns from one of the largest coleus breeding populations in the world. C_LIO_LIWorking with this massive dataset, we are able to analyze quantitative relationships between maternal plants and their progeny, identify features that underlie breeder-selections, and collect and compare consumer input on trait preferences. C_LIO_LIThis study is one of the most comprehensive explorations into complex color patterning in plant biology and provides new insights and tools for exploring the color pallet of the plant kingdom. C_LI

plant biology↗

bioRxiv2020

Directional phyllotactic bias in calatheas (Goeppertia, Marantaceae): a citizen science approach

Lateral organs arranged in spiral phyllotaxy are separated by the golden angle, {approx}137.5{degrees}, leading to handedness: either clockwise or counter-clockwise. In some species, leaves are asymmetric such that they are smaller and curved towards the side ascending the phyllotactic spiral. As such, these asymmetries lead to mirroring of leaf shapes in plants of opposite phyllotactic handedness. Previous reports had suggested that the pin-stripe calathea (Goeppertia ornata) may be exclusively of one phyllotactic direction, counter-clockwise, but had limited sampling to a single population. Here, we use a citizen science approach leveraging a social media poll, internet image searches, and in-person verification at nurseries in four countries and two continents to demonstrate that calatheas (Goeppertia spp.) around the world are biased towards counter-clockwise phyllotaxy. The possibility that this bias is genetic and its implications for models of phyllotaxy that assume handedness is stochastically specified in equal proportions is discussed.

plant biology↗

bioRxiv2020

Root system influence on high dimensional leaf phenotypes over the grapevine growing season

O_LIIn many perennial crops, grafting the root system of one individual to the shoot system of another individual has become an integral part of propagation performed at industrial scales to enhance pest, disease, and stress tolerance and to regulate yield and vigor. Grafted plants offer important experimental systems for understanding the extent and seasonality of root system effects on shoot system biology. C_LIO_LIUsing an experimental vineyard where a common scion Chambourcin is growing ungrafted and grafted to three different rootstocks, we explore associations between root system genotype and leaf phenotypes in grafted grapevines across a growing season. We quantified five high-dimensional leaf phenotyping modalities: ionomics, metabolomics, transcriptomics, morphometrics, and physiology and show that rootstock influence is subtle but ubiquitous across modalities. C_LIO_LIWe find strong signatures of rootstock influence on the leaf ionome, with unique signatures detected at each phenological stage. Moreover, all phenotypes and patterns of phenotypic covariation were highly dynamic across the season. C_LIO_LIThese findings expand upon previously identified patterns to suggest that the influence of root system on shoot system phenotypes is complex and broad understanding necessitates volumes of high-dimensional, multi-scale data previously unmet. C_LI

plant biology↗

bioRxiv2020

Composite modeling of leaf shape across shoots discriminates Vitis species better than individual leaves

Premise of studyLeaf morphology is dynamic, continuously deforming during leaf expansion and among leaves within a shoot. We measured leaf morphology from over 200 vines over four years, and modeled changes in leaf shape along the shoot to determine if a composite "shape of shapes" can better capture variation and predict species identity compared to individual leaves. MethodsUsing homologous universal landmarks found in grapevine leaves, we modeled various morphological features as a polynomial function of leaf node. The resulting functions are used to reconstruct modeled leaf shapes across shoots, generating composite leaves that comprehensively capture the spectrum of possible leaf morphologies. 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 species identity of previously unassigned vines, 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, as well as serving as a better predictor of species identity than individual leaves.

plant biology↗

bioRxiv2020

Vein-to-blade ratio is an allometric indicator of climate-induced changes in grapevine leaf size and shape

PremiseAs a leaf expands, its shape dynamically changes. Previously, we documented an allometric relationship between vein and blade area in grapevine leaves. Larger leaves have a smaller ratio of primary and secondary vein area relative to blade area compared to smaller leaves. We sought to use allometry as an indicator of leaf size to measure the environmental effects of climate on grapevine leaf morphology. MethodsWe measure the ratio of vein-to-blade area in 8,412 leaves from the same 208 vines across four growing seasons (2013, 2015, 2016, and 2017) using 21 homologous landmarks. Matching leaves by vine and node, we correlate size and shape of grapevine leaves with climate variables. Key resultsVein-to-blade ratio varies strongly between years in ways that blade or vein area do not. Maximum daily temperature and to a lesser degree precipitation are the most strongly correlated climate variables with vein-to-blade ratio, indicating that smaller leaves are associated with heat waves and drought. Leaf count and overall leaf area of shoots and the vineyard population studied also diminish with heat and drought. Grapevine leaf primordia initiate in buds the year prior to when they emerge, and we find that climate during the previous growing season exerts the largest statistical effects over these relationships. ConclusionsOur results demonstrate the profound effects of heat and drought on the vegetative morphology of grapevines and show that vein-to-blade ratio is a strong allometric indicator of the effects of climate on grapevine leaf morphology.

plant biology↗

bioRxiv2020

The shapes of wine and table grape leaves: an ampelometric study inspired by the methods of Pierre Galet

The shapes of grapevine leaves have been critical to correctly identify economically important varieties throughout history. The correspondence of homologous features in nearly all grapevine species and varieties has enabled advanced morphometric approaches to mathematically classify leaf shape. These approaches either model leaves through the measurement of numerous vein lengths and angles or measure a finite number of corresponding landmarks and use Procrustean approaches to superimpose points and perform statistical analyses. Hand illustrations, too, play an important role in grapevine identification, as details omitted using the above methods can be visualized. Here, I use a saturating number of pseudo-landmarks to capture intricate, local features in grapevine leaves: the curvature of veins and the shapes of serrations. Using these points, averaged leaf shapes for 60 varieties of wine and table grapes are calculated that preserve features. A pairwise Procrustes distance matrix of the overall morphological similarity of each variety to the other classifies leaves into two main groups--deeply lobed and more entire--that correspond to the measurements of sinus depth by Pierre Galet. Using the system of Galet, pseudo-landmarks are converted into relative distance and angle measurements. Both Galet-inspired and Procrustean methods allow increased accuracy in predicting variety compared to a finite number of landmarks. Using Procrustean pseudo-landmarks captures grapevine leaf shape at the same level of detail as drawings and provides a quantitative method to arrive at mean leaf shapes representing varieties that can be used within a predictive statistical framework.

plant biology↗

bioRxiv2019

Rootstock choice can dramatically affect grapevine growth

Grape growers use rootstocks to provide protection against pests and pathogens and to modulate viticulture performance such as shoot growth. Our study examined two grapevine scion varieties ( Chardonnay and Cabernet Sauvignon) grafted to 15 different rootstocks and determined the effect of rootstocks on eight traits important to viticulture. We assessed the vines across five years and identified both year and variety as contributing strongly to trait variation. The effect of rootstock was relatively consistent across years and varieties, explaining between 8.99% and 9.78% of the variation in growth-related traits including yield, pruning weight, berry weight, and Ravaz index (yield to pruning weight ratio). Increases in yield due to rootstock were generally the result of increases in berry weight, likely due to increased water uptake by vines grafted to a particular rootstock. We demonstrated a greater than 50% increase in yield, pruning weight, or Ravaz index by choosing the optimal rootstock, indicating that rootstock choice is crucial for grape growers looking to improve vine performance.

plant biology↗

bioRxiv2019

Rapid leaf trait response to growing-season meteorology in Vitis: Implications for leaf physiognomic paleoclimate reconstructions

PREMISE OF THE STUDYThe size and shape (physiognomy) of woody, dicotyledonous angiosperm leaves are correlated with climate and these relationships have been used to develop. proxies. These proxies assume that leaf morphology plastically responds to meteorological conditions and that leaf traits change isometrically through development.\n\nMETHODSWe used Digital Leaf Physiognomy (DiLP) to measure leaf characters of multiple Vitis species from the USDA Germplasm Repository in Geneva, NY from the 2012-2013 and 2014-2015 growing seasons. These growing seasons had different temperature and precipitation.\n\nKEY RESULTSWe found three primary results: (1) there were predictable significant differences in leaf characters in leaves of different developmental stages along the vine, (2) there were significant differences in leaf characters in leaves of the same developmental stage between the growing seasons, and (3) there were significant differences in leaf characters between growing seasons.\n\nCONCLUSIONSWe found that Vitis leaf shape had the strongest relationship with growing season meteorological conditions in taxa growing in their native range. In addition, leaves have variable phenotypic plasticity along the vine. We interpret that the meteorological signal was strongest in those leaves that have completed allometric expansion. This is significant for leaf physiognomic-paleoclimate proxies because these leaves are most likely to be preserved in leaf litter and reflect the type of leaves included in paleoclimate reconstructions. We found that leaf development does have the potential to be a confounding factor, but it is unlikely to exert a significant influence on analysis due to differential preservation potential.

plant biology↗

bioRxiv2019

Leaf shape is a predictor of fruit quality and cultivar performance in tomato

O_LICommercial tomato (Solanum lycopersicum) is one of the most widely grown vegetable crops worldwide. Heirloom tomatoes retain extensive genetic diversity and a considerable range of fruit quality and leaf morphological traits.\nC_LIO_LIHere the role of leaf morphology was investigated for its impact on fruit quality. Heirloom cultivars were grown in field conditions and BRIX by Yield (BY) and other traits measured over a fourteen-week period. The complex relationships among these morphological and physiological traits were evaluated using PLS-Path Modeling, and a consensus model developed.\nC_LIO_LIPhotosynthesis contributed strongly to vegetative biomass and sugar content of fruits but had a negative impact on yield. Conversely leaf shape, specifically rounder leaves, had a strong positive impact on both fruit sugar content and yield. Cultivars such as Stupice and Glacier, with very round leaves, had the highest performance in both fruit sugar and yield. Our model accurately predicted BY for two commercial cultivars using leaf shape data as input.\nC_LIO_LIThis study revealed the importance of leaf shape to fruit quality in tomato, with rounder leaves having significantly improved fruit quality. This correlation was maintained across a range of diverse genetic backgrounds and shows the importance of leaf morphology in tomato crop improvement.\nC_LI

plant biology↗

bioRxiv2018

Rootstock effects on scion phenotypes in a ‘Chambourcin’ experimental vineyard

Understanding how root systems modulate shoot system phenotypes is a fundamental question in plant biology and will be useful in developing resilient agricultural crops. Grafting is a common horticultural practice that joins the roots (rootstock) of one plant to the shoot (scion) of another, providing an excellent method for investigating how these two organ systems affect each other. In this study, we use the French-American hybrid grapevine Chambourcin (Vitis L.) as a model to explore the rootstock-scion relationship. We examined leaf shape, ion concentrations, and gene expression in Chambourcin grown own-rooted as well as grafted to three different rootstocks ( SO4, 1103P and 3309C) across two years and three different irrigation treatments. Results described here demonstrate that 1) the largest source of variation in leaf shape stems from the interaction of rootstock by irrigation; 2) leaf position, but also rootstock and rootstock by irrigation interaction, are the primary sources of variation in leaf ion concentrations; and 3) gene expression in scion leaves exhibited significantly different patterns of gene expression from ungrafted vines, and these expression patterns were rootstock-specific. Our work provides an initial description of the subtle and complex effect of grafting on Chambourcin leaf morphology, ionomics and gene expression in grapevine scions. Further work across multiple years, environments and additional phenotypes is required in order to determine how the relationship between the rootstock and the scion can best be leveraged for adapting grapevines to a changing climate.

plant biology↗

bioRxiv2017

Bundle sheath extensions in tomato affect leaf phenotypic plasticity in response to irradiance

Coordination between structural and physiological traits is key to plants responses to environmental fluctuations. In heterobaric leaves, bundle sheath extensions (BSEs) increase photosynthetic performance (light-saturated rates of photosynthesis, Amax) and water transport capacity (leaf hydraulic conductance, Kleaf). However, it is not clear how BSEs affect these and other leaf developmental and physiological parameters in response to environmental conditions. The obscuravenosa (obv) mutation, found in many commercial tomato varieties, leads to absence of BSEs. We examined structural and physiological traits of tomato heterobaric and homobaric (obv) near-isogenic lines (NILs) grown at two different irradiance levels. Kleaf, minor vein density and stomatal pore area index decreased with shading in heterobaric but not in homobaric leaves, which show similarly lower values in both conditions. Homobaric plants, on the other hand, showed increased Amax, leaf intercellular air spaces and mesophyll surface area exposed to intercellular airspace (Smes) in comparison with heterobaric plants when both were grown in the shade. BSEs further affected carbon isotope discrimination, a proxy for long-term water-use efficiency. BSEs confer plasticity in traits related to leaf structure and function in response to irradiance levels and might act as a hub integrating leaf structure, photosynthetic function and water supply and demand.\n\nSummary statementThe presence of bundle sheath extension (BSEs) defines leaves as heterobaric, as opposed to homobaric leaves that lack them. Multiple functions have been proposed for BSEs, but their impact on different environmental conditions is still unclear. Here, we compared a tomato (Solanum lycopersicum) homobaric mutant lacking BSEs with its corresponding heterobaric wild-type, grown under two irradiance conditions. We show that the presence of BSEs differentially alters various physiological and anatomical parameters in response to growth irradiance. We propose that BSEs could act as hubs coordinating leaf plasticity in response to environmental factors.\n\nArticle typeResearch article

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bioRxiv2017

Persistent homology demarcates a leaf morphospace

Current morphometric methods that comprehensively measure shape cannot compare the disparate leaf shapes found in seed plants and are sensitive to processing artifacts. We explore the use of persistent homology, a topological method applied across the scales of a function, to overcome these limitations. The described method isolates subsets of shape features and measures the spatial relationship of neighboring pixel densities in a shape. We apply the method to the analysis of 182,707 leaves, both published and unpublished, representing 141 plant families collected from 75 sites throughout the world. By measuring leaves from throughout the seed plants using persistent homology, a defined morphospace comparing all leaves is demarcated. Clear differences in shape between major phylogenetic groups are detected and estimates of leaf shape diversity within plant families are made. This approach does not only predict plant family, but also the collection site, confirming phylogenetically invariant morphological features that characterize leaves from specific locations. The application of a persistent homology method to measure leaf shape allows for a unified morphometric framework to measure plant form, including shape and branching architectures.

plant biology↗

bioRxiv2017

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.

plant biology↗

bioRxiv2017

Genetic Architecture and Molecular Networks Underlying Leaf Thickness in Desert-Adapted Tomato Solanum pennellii

Thicker leaves allow plants to grow in water-limited conditions. However, our understanding of the genetic underpinnings of this highly functional leaf shape trait is poor. We used a custom-built confocal profilometer to directly measure leaf thickness in a set of introgression lines (ILs) derived from the desert tomato species Solanum pennellii, and identified quantitative trait loci (QTL). We report evidence of a complex genetic architecture of this trait and roles for both genetic and environmental factors. Several ILs with thick leaves have dramatically elongated palisade mesophyll cells and, in some cases, increased leaf ploidy. We characterized thick ILs 2-5 and 4-3 in detail and found increased mesophyll cell size and leaf ploidy levels, suggesting that endoreduplication underpins leaf thickness in tomato. Next, we queried the transcriptomes and inferred Dynamic Bayesian Networks of gene expression across early leaf ontogeny in these lines to compare the molecular networks that pattern leaf thickness. We show that thick ILs share S. pennellii-like expression profiles for putative regulators of cell shape and meristem determinacy, as well as a general signature of cell cycle related gene expression. However, our network data suggest that leaf thickness in these two lines is patterned by at least partially distinct mechanisms. Consistent with this hypothesis, double homozygote lines combining introgression segments from these two ILs show additive phenotypes including thick leaves, higher ploidy levels and larger palisade mesophyll cells. Collectively, these data establish a framework of genetic, anatomical, and molecular mechanisms that pattern leaf thickness in desert-adapted tomato.

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