A genotype-specific architectural and physiological profile is involved in the flowering regularity of apple trees.

In polycarpic plants, meristem fate varies within individuals in a given year. In perennials, the proportion of floral induction (FI) in meristems also varies between consecutive years and among genotypes of a given species. Previous studies have suggested that FI of meristems could be determined by the within-plant competition for carbohydrates and by hormone signaling as key components of the flowering pathway. At the genotypic level, variability in FI was also associated with variability in architectural traits. However, the part of genotype-dependent variability in FI that can be explained by either tree architecture or tree physiology is still not fully understood. This study aimed at deciphering the respective effect of architectural and physiological traits on FI variability within apple trees by comparing six genotypes with contrasted architectures. Shoot type demography as well as the flowering and fruit production patterns were followed over 6 years and characterized by different indexes. Architectural morphotypes were then defined based on architectural traits using a clustering approach. For two successive years, non-structural starch content in leaf, stem and meristems, and hormonal contents (gibberellins, cytokinins, auxin and abscisic acid) in meristems were quantified and correlated to FI within-tree proportions. Based on a multi-step regression analysis, cytokinins and gibberellins content in meristem, starch content in leaves and the proportion of long shoots in tree annual growth were shown to contribute to FI. Although the predictive linear model of FI was common to all genotypes, each of the explicative variables had a different weight in FI determination, depending on the genotype. Our results therefore suggest both a common determination model and a genotype-specific architectural and physiological profile linked to its flowering behavior.

Tree architecture, light interception and water-use related traits are controlled by different genomic regions in an apple tree core collection.

Tree architecture shows large genotypic variability, but how this affects water-deficit responses is poorly understood. To assess the possibility of reaching ideotypes with adequate combinations of architectural and functional traits in the face of climate change, we combined high-throughput field phenotyping and genome-wide association studies (GWAS) on an apple tree (Malus domestica) core-collection. We used terrestrial light detection and ranging (T-LiDAR) scanning and airborne multispectral and thermal imagery to monitor tree architecture, canopy shape, light interception, vegetation indices and transpiration on 241 apple cultivars submitted to progressive field soil drying. GWAS was performed with single nucleotide polymorphism (SNP)-by-SNP and multi-SNP methods. Large phenotypic and genetic variability was observed for all traits examined within the collection, especially canopy surface temperature in both well-watered and water deficit conditions, suggesting control of water loss was largely genotype-dependent. Robust genomic associations revealed independent genetic control for the architectural and functional traits. Screening associated genomic regions revealed candidate genes involved in relevant pathways for each trait. We show that multiple allelic combinations exist for all studied traits within this collection. This opens promising avenues to jointly optimize tree architecture, light interception and water use in breeding strategies. Genotypes carrying favourable alleles depending on environmental scenarios and production objectives could thus be targeted.

Spatial variability in carbon- and nitrogen-related traits in apple trees: the effects of the light environment and crop load.

Photosynthetic carbon assimilation rates are highly dependent on environmental factors such as light availability and on metabolic limitations such as the demand for carbon by sink organs. The relative effects of light and sink demand on photosynthesis in perennial plants such as trees remain poorly characterized. The aim of the present study was therefore to characterize the relationships between light and fruit load on a range of leaf traits including photosynthesis, non-structural carbohydrate content, leaf structure, and nitrogen-related variables in fruiting ('ON') and non-fruiting ('OFF') 'Golden Delicious' apple trees. We show that crop status (at the tree scale) exerts a greater influence over leaf traits than the local light environment or the local fruit load. High rates of photosynthesis were observed in the ON trees. This was correlated with a high leaf nitrogen content. In contrast, little spatial variability in photosynthesis rates was observed in the OFF trees. The lack of variation in photosynthesis rates was associated with high leaf non-structural carbohydrate content at the tree level. Taken together, these results suggest that low carbon demand leads to feedback limitation on photosynthesis resulting in a low level of within-tree variability. These findings provide new insights into carbon and nitrogen allocations within trees, which are heavily dependent on carbon demand.

A Comparative Study on the Branching Pattern of Monocyclic and Bicyclic Shoots of Apple cv. "Fuji".

The development of tree architecture results from shoot growth and branching, but their relationship is still not fully understood. The goal of this study was to determine the effect of parent shoot growth characteristics on branching patterns in terms of polycyclism, growth duration (GD), and growth period (GP), considering apple tree as a case study. Weekly shoot growth records were collected from 227 shoots during their second year of growth and the resulting branching patterns from the following year. The branching patterns were compared between the different shoot categories, using hidden semi-Markov models. Our results showed that the branching pattern was similar in bicyclic and monocyclic shoots with a long GD. The number of floral laterals, and the frequency and length of the floral zones, increased with GD. Moreover, a long GD led to strong acrotony, due to the high occurrence of a vegetative zone with long laterals in the distal position of the shoot. In bicyclic shoots, an early GP of the second GU led to more frequent and longer floral zones than a late GP. Therefore, the GD was the strongest driver of the branching pattern, and GP modulated the flowering capacity. The main similarities among shoot categories resulted from the existence of latent buds and floral zones associated with growth cessation periods. Even though flowering was more abundant during the early GP, the positions of floral zones indicated that induction in axillary meristems can also occur late in the season. This study provides new knowledge regarding the relationships between the dynamics of parent shoot growth and axillary meristem fates, with key consequences on flowering abundance and positions.

Modelling transport of inhibiting and activating signals and their combined effects on floral induction: application to apple tree.

Floral induction (FI) in shoot apical meristems (SAM) is assumed to be triggered by antagonistic endogenous signals. In fruit trees, FI occurs in some SAM only and is determined by activating and inhibiting signals originating from leaves and fruit, respectively. We developed a model (SigFlow) to quantify on 3D structures the combined impact of such signals and distances at which they act on SAM. Signal transport was simulated considering a signal 'attenuation' parameter, whereas SAM fate was determined by probability functions depending on signal quantities. Model behaviour was assessed on simple structures before being calibrated and validated on a unique experimental dataset of 3D digitized apple trees with contrasted crop loads and subjected to leaf and fruit removal at different scales of tree organization. Model parameter estimations and comparisons of two signal combination functions led us to formulate new assumptions on the mechanisms involved: (i) the activating signal could be transported at shorter distances than the inhibiting one (roughly 50 cm vs 1 m) (ii) both signals jointly act to determine FI with SAM being more sensitive to inhibiting signal than activating one. Finally, the genericity of the model is promising to further understand the physiological and architectural determinisms of FI in plants.

Environmental and trophic determinism of fruit abscission and outlook with climate change in tropical regions.

Fruit abscission facilitates the optimal conditions and timing of seed dispersal. Environmental regulation of tropical fruit abscission has received little attention, even though climate change may have its strongest impacts in tropical regions. In this study, oil palm fruit abscission was monitored during multiple years in the Benin Republic to take advantage of the climatic seasonality and the continuous fruit production by this species. An innovative multivariable statistical method was used to identify the best predictors of fruit abscission among a set of climate and ecophysiological variables, and the stage of inflorescence and fruit bunch development when the variables are perceived. The effects of climate scenarios on fruit abscission were then predicted based on the calibrated model. We found complex regulation takes place at specific stages of inflorescence and bunch development, even long before the fruit abscission zone is competent to execute abscission. Among the predictors selected, temperature variations during inflorescence and fruit bunch development are major determinants of the fruit abscission process. Furthermore, the timing of ripe fruit drop is determined by temperature in combination with the trophic status. Finally, climate simulations revealed that the abscission process is robust and is more affected by seasonal variations than by extreme scenarios. Our investigations highlighted the central function of the abscission zone as the sensor of environmental signals during reproductive development. Coupling ecophysiological and statistical modeling was an efficient approach to disentangle this complex environmental regulation.

Impact of Within-Tree Organ Distances on Floral Induction and Fruit Growth in Apple Tree: Implication of Carbohydrate and Gibberellin Organ Contents.

In plants, organs are inter-dependent for growth and development. Here, we aimed to investigate the distance at which interaction between organs operates and the relative contribution of within-tree variation in carbohydrate and hormonal contents on floral induction and fruit growth, in a fruit tree case study. Manipulations of leaf and fruit numbers were performed in two years on "Golden delicious" apple trees, at the shoot or branch scale or one side of Y-shape trees. For each treatment, floral induction proportion and mean fruit weight were recorded. Gibberellins content in shoot apical meristems, photosynthesis, and non-structural carbohydrate concentrations in organs were measured. Floral induction was promoted by leaf presence and fruit absence but was not associated with non-structural content in meristems. This suggests a combined action of promoting and inhibiting signals originating from leaves and fruit, and involving gibberellins. Nevertheless, these signals act at short distance only since leaf or fruit presence at long distances had no effect on floral induction. Conversely, fruit growth was affected by leaf presence even at long distances when sink demands were imbalanced within the tree, suggesting long distance transport of carbohydrates. We thus clarified the inter-dependence and distance effect among organs, therefore their degree of autonomy that appeared dependent on the process considered, floral induction or fruit growth.

Multi-scale high-throughput phenotyping of apple architectural and functional traits in orchard reveals genotypic variability under contrasted watering regimes.

Despite previous reports on the genotypic variation of architectural and functional traits in fruit trees, phenotyping large populations in the field remains challenging. In this study, we used high-throughput phenotyping methods on an apple tree core-collection (1000 individuals) grown under contrasted watering regimes. First, architectural phenotyping was achieved using T-LiDAR scans for estimating convex and alpha hull volumes and the silhouette to total leaf area ratio (STAR). Second, a semi-empirical index (I PL) was computed from chlorophyll fluorescence measurements, as a proxy for leaf photosynthesis. Last, thermal infrared and multispectral airborne imaging was used for computing canopy temperature variations, water deficit, and vegetation indices. All traits estimated by these methods were compared to low-throughput in planta measurements. Vegetation indices and alpha hull volumes were significantly correlated with tree leaf area and trunk cross sectional area, while I PL values showed strong correlations with photosynthesis measurements collected on an independent leaf dataset. By contrast, correlations between stomatal conductance and canopy temperature estimated from airborne images were lower, emphasizing discrepancies across measurement scales. High heritability values were obtained for almost all the traits except leaf photosynthesis, likely due to large intra-tree variation. Genotypic means were used in a clustering procedure that defined six classes of architectural and functional combinations. Differences between groups showed several combinations between architectural and functional traits, suggesting independent genetic controls. This study demonstrates the feasibility and relevance of combining multi-scale high-throughput methods and paves the way to explore the genetic bases of architectural and functional variations in woody crops in field conditions.

Neoformation and summer arrest are common sources of tree plasticity in response to water stress of apple cultivars.

BACKGROUND AND AIMS: Depending on the species, water stress affects different growth and developmental processes, mainly due to changes in hydraulic properties and hormonal signalling. This study compared the impact of water stress on tree development and organ growth in three apple cultivars. METHODS: Trees were differentially irrigated to induce water stress or to provide well-watered conditions in their second and third years of growth. Effects of water stress were evaluated at tree scale by shoot number and proportions of the different types of shoots, and at shoot scale by metamer appearance rate, growth duration and arrest time, as well as organ size. KEY RESULTS: Water stress promoted early growth cessation, prolonged summer arrests and decreased growth resumptions, thus modifying within-tree shoot demography in favour of short shoots. Growth cessations occurred in mild water stress conditions before any difference in stem water potential appeared. No major impact was observed on organ size. Consistently with tree ontogeny, the number of shoots that resumed growth after summer arrest decreased with years, but more in water-stressed than well-watered conditions. CONCLUSIONS: Even though the impact of water stress differed slightly among cultivars, the reduction in neoformation and increase in summer arrest played a common role in apple tree morphological responses and led to stress avoidance by early reduction of tree leaf area.

Long proleptic and sylleptic shoots in peach (Prunus persica L. Batsch) trees have similar, predetermined, maximum numbers of nodes and bud fate patterns.

BACKGROUND AND AIMS: In peach (Prunus persica) trees, three types of shoots can be distinguished depending on the time of their appearance: sylleptic, proleptic and epicormic. On proleptic shoots, an average of ten phytomers are preformed in dormant buds prior to shoot growth after bud-break, whereas all phytomers are considered neoformed in sylleptic and epicormic shoots. However, casual observations indicated that proleptic and sylleptic shoots appear quite similar in number of phytomers and structure in spite of their different origins. The goal of this research was to test the hypothesis that both proleptic and sylleptic shoots exhibit similar growth characteristics by analysing their node numbers and bud fate patterns. If their growth characteristics are similar, it would indicate that the structure of both types of shoots is primarily under genetic rather than environmental control. METHODS: The number of phytomers and bud fate patterns of proleptic and sylleptic shoots of four peach cultivars grown in the same location (Winters, California) were analysed and characterized using hidden semi-Markov models. Field data were collected during winter 2016, just prior to floral bud-break. KEY RESULTS: Sylleptic shoots tended to have slightly fewer phytomers than proleptic shoots of the same cultivars. The bud fate patterns along proleptic and sylleptic shoots were remarkably similar for all the cultivars, although proleptic shoots started growing earlier (at least 1 month) in the spring than sylleptic shoots. CONCLUSIONS: This study provides strong evidence for the semi-deterministic nature of both proleptic and sylleptic shoots across four peach cultivars in terms of number of phytomers and bud fate patterns along shoots. It is apparent that the overall structure of shoots with similar numbers of phytomers was under similar genetic control for the two shoot types. Understanding shoot structural characteristics can aid in phenotypic characterization of vegetative growth of trees and in providing a foundation for vegetative management of fruit trees in horticultural settings.

Growth and carbon balance are differently regulated by tree and shoot fruiting contexts: an integrative study on apple genotypes with contrasted bearing patterns.

In plants, carbon source-sink relationships are assumed to affect their reproductive effort. In fruit trees, carbon source-sink relationships are likely to be involved in their fruiting behavior. In apple, a large variability in fruiting behaviors exists, from regular to biennial, which has been related to the within-tree synchronization vs desynchronization of floral induction in buds. In this study, we analyzed if carbon assimilation, availability and fluxes as well as shoot growth differ in apple genotypes with contrasted behaviors. Another aim was to determine the scale of plant organization at which growth and carbon balance are regulated. The study was carried out on 16 genotypes belonging to three classes: (i) biennial, (ii) regular with a high production of floral buds every year and (iii) regular, displaying desynchronized bud fates in each year. Three shoot categories, vegetative and reproductive shoots with or without fruits, were included. This study shows that shoot growth and carbon balance are differentially regulated by tree and shoot fruiting contexts. Shoot growth was determined by the shoot fruiting context, or by the type of shoot itself, since vegetative shoots were always longer than reproductive shoots whatever the tree crop load. Leaf photosynthesis depended on the tree crop load only, irrespective of the shoot category or the genotypic class. Starch content was also strongly affected by the tree crop load with some adjustments of the carbon balance among shoots since starch content was lower, at least at some dates, in shoots with fruits compared with the shoots without fruits within the same trees. Finally, the genotypic differences in terms of shoot carbon balance partly matched with genotypic bearing patterns. Nevertheless, carbon content in buds and the role of gibberellins produced by seeds as well as the distances at which they could affect floral induction should be further analyzed.

Designing oil palm architectural ideotypes for optimal light interception and carbon assimilation through a sensitivity analysis of leaf traits.

Background and Aims: Enhancement of light harvesting in annual crops has successfully led to yield increases since the green revolution. Such an improvement has mainly been achieved by selecting plants with optimal canopy architecture for specific agronomic practices. For perennials such as oil palm, breeding programmes were focused more on fruit yield, but now aim at exploring more complex traits. The aim of the present study is to investigate potential improvements in light interception and carbon assimilation in the study case of oil palm, by manipulating leaf traits and proposing architectural ideotypes. Methods: Sensitivity analyses (Morris method and metamodel) were performed on a functional-structural plant model recently developed for oil palm which takes into account genetic variability, in order to virtually assess the impact of plant architecture on light interception efficiency and potential carbon acquisition. Key Results: The most sensitive parameters found over plant development were those related to leaf area (rachis length, number of leaflets, leaflet morphology), although fine attributes related to leaf geometry showed increasing influence when the canopy became closed. In adult stands, optimized carbon assimilation was estimated on plants with a leaf area index between 3.2 and 5.5 m2 m-2 (corresponding to usual agronomic conditions), with erect leaves, short rachis and petiole, and high number of leaflets on the rachis. Four architectural ideotypes for carbon assimilation are proposed based on specific combinations of organ dimensions and arrangement that limit mutual shading and optimize light distribution within the plant crown. Conclusions: A rapid set-up of leaf area is critical at young age to optimize light interception and subsequently carbon acquisition. At the adult stage, optimization of carbon assimilation could be achieved through specific combinations of architectural traits. The proposition of multiple morphotypes with comparable level of carbon assimilation opens the way to further investigate ideotypes carrying an optimal trade-off between carbon assimilation, plant transpiration and biomass partitioning.

The impact of long-term water stress on tree architecture and production is related to changes in transitions between vegetative and reproductive growth in the 'Granny Smith' apple cultivar.

Water stress (WS) generates a number of physiological and morphological responses in plants that depend on the intensity and duration of stress as well as the plant species and development stage. In perennial plants, WS may affect plant development through cumulative effects that modify plant functions, architecture and production over time. Plant architecture depends on the fate of the terminal and axillary buds that can give rise, in the particular case of apple, to reproductive or vegetative growth units (GUs) of different lengths. In this study, the impact of long-term WS (7 years) on the fate of terminal and axillary buds was investigated in relation to flowering occurrence and production pattern (biennial vs regular) in the 'Granny Smith' cultivar. It was observed that WS decreased the total number of GUs per branch, regardless of their type. Conversely, WS did not modify the timing of the two successive developmental phases characterized by the production of long and medium GUs and an alternation of floral GUs over time, respectively. The analysis of GU successions over time using a variable-order Markov chain that included both the effects of the predecessor and water treatment revealed that WS reduced the transition towards long and medium GUs and increased the transition toward floral, short and dead GUs. WS also slightly increased the proportion of axillary floral GUs. The higher relative frequency of floral GUs compared with vegetative ones reduced the tendency to biennial bearing under WS. The accelerated ontogenetic trend observed under WS suggests lower vegetative growth that could, in turn, be beneficial to floral induction and fruit set.

Integrating mixed-effect models into an architectural plant model to simulate inter- and intra-progeny variability: a case study on oil palm (Elaeis guineensis Jacq.).

Three-dimensional (3D) reconstruction of plants is time-consuming and involves considerable levels of data acquisition. This is possibly one reason why the integration of genetic variability into 3D architectural models has so far been largely overlooked. In this study, an allometry-based approach was developed to account for architectural variability in 3D architectural models of oil palm (Elaeis guineensis Jacq.) as a case study. Allometric relationships were used to model architectural traits from individual leaflets to the entire crown while accounting for ontogenetic and morphogenetic gradients. Inter- and intra-progeny variabilities were evaluated for each trait and mixed-effect models were used to estimate the mean and variance parameters required for complete 3D virtual plants. Significant differences in leaf geometry (petiole length, density of leaflets, and rachis curvature) and leaflet morphology (gradients of leaflet length and width) were detected between and within progenies and were modelled in order to generate populations of plants that were consistent with the observed populations. The application of mixed-effect models on allometric relationships highlighted an interesting trade-off between model accuracy and ease of defining parameters for the 3D reconstruction of plants while at the same time integrating their observed variability. Future research will be dedicated to sensitivity analyses coupling the structural model presented here with a radiative balance model in order to identify the key architectural traits involved in light interception efficiency.

Simulation of carbon allocation and organ growth variability in apple tree by connecting architectural and source-sink models.

BACKGROUND AND AIMS: Plant growth depends on carbon availability and allocation among organs. QualiTree has been designed to simulate carbon allocation and partitioning in the peach tree (Prunus persica), whereas MappleT is dedicated to the simulation of apple tree (Malus × domestica) architecture. The objective of this study was to couple both models and adapt QualiTree to apple trees to simulate organ growth traits and their within-tree variability. METHODS: MappleT was used to generate architectures corresponding to the 'Fuji' cultivar, accounting for the variability within and among individuals. These architectures were input into QualiTree to simulate shoot and fruit growth during a growth cycle. We modified QualiTree to account for the observed shoot polymorphism in apple trees, i.e. different classes (long, medium and short) that were characterized by different growth function parameters. Model outputs were compared with observed 3D tree geometries, considering shoot and final fruit size and growth dynamics. KEY RESULTS: The modelling approach connecting MappleT and QualiTree was appropriate to the simulation of growth and architectural characteristics at the tree scale (plant leaf area, shoot number and types, fruit weight at harvest). At the shoot scale, mean fruit weight and its variability within trees was accurately simulated, whereas the model tended to overestimate individual shoot leaf area and underestimate its variability for each shoot type. Varying the parameter related to the intensity of carbon exchange between shoots revealed that behaviour intermediate between shoot autonomy and a common assimilate pool was required to properly simulate within-tree fruit growth variability. Moreover, the model correctly dealt with the crop load effect on organ growth. CONCLUSIONS: This study provides understanding of the integration of shoot ontogenetic properties, carbon supply and transport between entities for simulating organ growth in trees. Further improvements regarding the integration of retroaction loops between carbon allocation and the resulting plant architecture are expected to allow multi-year simulations.

Combining Genome-Wide Information with a Functional Structural Plant Model to Simulate 1-Year-Old Apple Tree Architecture.

In crops, optimizing target traits in breeding programs can be fostered by selecting appropriate combinations of architectural traits which determine light interception and carbon acquisition. In apple tree, architectural traits were observed to be under genetic control. However, architectural traits also result from many organogenetic and morphological processes interacting with the environment. The present study aimed at combining a FSPM built for apple tree, MAppleT, with genetic determinisms of architectural traits, previously described in a bi-parental population. We focused on parameters related to organogenesis (phyllochron and immediate branching) and morphogenesis processes (internode length and leaf area) during the first year of tree growth. Two independent datasets collected in 2004 and 2007 on 116 genotypes, issued from a 'Starkrimson' × 'Granny Smith' cross, were used. The phyllochron was estimated as a function of thermal time and sylleptic branching was modeled subsequently depending on phyllochron. From a genetic map built with SNPs, marker effects were estimated on four MAppleT parameters with rrBLUP, using 2007 data. These effects were then considered in MAppleT to simulate tree development in the two climatic conditions. The genome wide prediction model gave consistent estimations of parameter values with correlation coefficients between observed values and estimated values from SNP markers ranging from 0.79 to 0.96. However, the accuracy of the prediction model following cross validation schemas was lower. Three integrative traits (the number of leaves, trunk length, and number of sylleptic laterals) were considered for validating MAppleT simulations. In 2007 climatic conditions, simulated values were close to observations, highlighting the correct simulation of genetic variability. However, in 2004 conditions which were not used for model calibration, the simulations differed from observations. This study demonstrates the possibility of integrating genome-based information in a FSPM for a perennial fruit tree. It also showed that further improvements are required for improving the prediction ability. Especially temperature effect should be extended and other factors taken into account for modeling GxE interactions. Improvements could also be expected by considering larger populations and by testing other genome wide prediction models. Despite these limitations, this study opens new possibilities for supporting plant breeding by in silico evaluations of the impact of genotypic polymorphisms on plant integrative phenotypes.

Genetic Variation of Morphological Traits and Transpiration in an Apple Core Collection under Well-Watered Conditions: Towards the Identification of Morphotypes with High Water Use Efficiency.

Water use efficiency (WUE) is a quantitative measurement which improvement is a major issue in the context of global warming and restrictions in water availability for agriculture. In this study, we aimed at studying the variation and genetic control of WUE and the respective role of its components (plant biomass and transpiration) in a perennial fruit crop. We explored an INRA apple core collection grown in a phenotyping platform to screen one-year-old scions for their accumulated biomass, transpiration and WUE under optimal growing conditions. Plant biomass was decompose into morphological components related to either growth or organ expansion. For each trait, nine mixed models were evaluated to account for the genetic effect and spatial heterogeneity inside the platform. The Best Linear Unbiased Predictors of genetic values were estimated after model selection. Mean broad-sense heritabilities were calculated from variance estimates. Heritability values indicated that biomass (0.76) and WUE (0.73) were under genetic control. This genetic control was lower in plant transpiration with an heritability of 0.54. Across the collection, biomass accounted for 70% of the WUE variability. A Hierarchical Ascendant Classification of the core collection indicated the existence of six groups of genotypes with contrasting morphology and WUE. Differences between morphotypes were interpreted as resulting from differences in the main processes responsible for plant growth: cell division leading to the generation of new organs and cell elongation leading to organ dimension. Although further studies will be necessary on mature trees with more complex architecture and multiple sinks such as fruits, this study is a first step for improving apple plant material for the use of water.

Relationships between biomass allocation, axis organogenesis and organ expansion under shading and water deficit conditions in grapevine.

The relationships between whole-plant growth and morphogenetic processes under abiotic stresses are still partly unknown. Whole-plant biomass growth can be decreased by many abiotic stresses, including water deficit and shading. Two experiments were performed on potted plants of one grapevine cultivar (Vitis vinifera L. cv. Syrah) subjected to watering and shading treatments. Under water stress, plants reduced their primary and secondary axis leaf production rate, whereas secondary axis budburst was relatively unaffected. Individual leaf area was reduced and a strong decrease in leaf expansion rate was observed. Under shading, primary axis organogenesis was maintained, both secondary axis budburst rate and phytomer appearance rate were decreased, and individual leaf area slightly increased. Specific leaf area did not change under soil water deficit, whereas it increased under shading. These results confirm the existence of dynamic changes in organ sink strength and biomass allocation patterns to favour plant leaf area growth under shading, and to reduce plant leaf area and water losses by transpiration under water stress. From a modelling point of view, this study shows that functional structural models based on a C balance are not fully relevant for simulating plant growth under abiotic constraints if they do not include non-trophic relationships (hormonal signalling or plant hydraulic properties) that modify organ sink strength according to abiotic constraints.

Using plant growth modeling to analyze C source-sink relations under drought: inter- and intraspecific comparison.

The ability to assimilate C and allocate non-structural carbohydrates (NSCs) to the most appropriate organs is crucial to maximize plant ecological or agronomic performance. Such C source and sink activities are differentially affected by environmental constraints. Under drought, plant growth is generally more sink than source limited as organ expansion or appearance rate is earlier and stronger affected than C assimilation. This favors plant survival and recovery but not always agronomic performance as NSC are stored rather than used for growth due to a modified metabolism in source and sink leaves. Such interactions between plant C and water balance are complex and plant modeling can help analyzing their impact on plant phenotype. This paper addresses the impact of trade-offs between C sink and source activities and plant production under drought, combining experimental and modeling approaches. Two contrasted monocotyledonous species (rice, oil palm) were studied. Experimentally, the sink limitation of plant growth under moderate drought was confirmed as well as the modifications in NSC metabolism in source and sink organs. Under severe stress, when C source became limiting, plant NSC concentration decreased. Two plant models dedicated to oil palm and rice morphogenesis were used to perform a sensitivity analysis and further explore how to optimize C sink and source drought sensitivity to maximize plant growth. Modeling results highlighted that optimal drought sensitivity depends both on drought type and species and that modeling is a great opportunity to analyze such complex processes. Further modeling needs and more generally the challenge of using models to support complex trait breeding are discussed.