Spatio-temporal analysis of strawberry architecture: insights into the control of branching and inflorescence complexity.

Plant architecture plays a major role in flowering and therefore in crop yield. Attempts to visualize and analyse strawberry plant architecture have been few to date. Here, we developed open-source software combining two- and three-dimensional representations of plant development over time along with statistical methods to explore the variability in spatio-temporal development of plant architecture in cultivated strawberry. We applied this software to six seasonal strawberry varieties whose plants were exhaustively described monthly at the node scale. Results showed that the architectural pattern of the strawberry plant is characterized by a decrease of the module complexity between the zeroth-order module (primary crown) and higher-order modules (lateral branch crowns and extension crowns). Furthermore, for each variety, we could identify traits with a central role in determining yield, such as date of appearance and number of branches. By modeling the spatial organization of axillary meristem fate on the zeroth-order module using a hidden hybrid Markov/semi-Markov mathematical model, we further identified three zones with different probabilities of production of branch crowns, dormant buds, or stolons. This open-source software will be of value to the scientific community and breeders in studying the influence of environmental and genetic cues on strawberry architecture and yield.

V-Mango: a functional-structural model of mango tree growth, development and fruit production.

BACKGROUND AND AIMS: Mango (Mangifera indica L.) is the fifth most widely produced fruit in the world. Its cultivation, mainly in tropical and sub-tropical regions, raises a number of issues such as the irregular fruit production across years, phenological asynchronisms that lead to long periods of pest and disease susceptibility, and the heterogeneity of fruit quality and maturity at harvest. To address these issues, we developed an integrative functional-structural plant model that synthesizes knowledge about the vegetative and reproductive development of the mango tree and opens up the possible simulation of cultivation practices. METHODS: We designed a model of architectural development in order to precisely characterize the intricate developmental processes of the mango tree. The appearance of botanical entities was decomposed into elementary stochastic events describing occurrence, intensity and timing of development. These events were determined by structural (position and fate of botanical entities) and temporal (appearance dates) factors. Daily growth and development of growth units and inflorescences were modelled using empirical distributions and thermal time. Fruit growth was determined using an ecophysiological model that simulated carbon- and water-related processes at the fruiting branch scale. KEY RESULTS: The model simulates the dynamics of the population of growth units, inflorescences and fruits at the tree scale during a growing cycle. Modelling the effects of structural and temporal factors makes it possible to simulate satisfactorily the complex interplays between vegetative and reproductive development. The model allowed the characterization of the susceptibility of mango tree to pests and the investigatation of the influence of tree architecture on fruit growth. CONCLUSIONS: This integrative functional-structural model simulates mango tree vegetative and reproductive development over successive growing cycles, allowing a precise characterization of tree phenology and fruit growth and production. The next step is to integrate the effects of cultivation practices, such as pruning, into the model.

Cytokinin signalling inhibitory fields provide robustness to phyllotaxis.

How biological systems generate reproducible patterns with high precision is a central question in science. The shoot apical meristem (SAM), a specialized tissue producing plant aerial organs, is a developmental system of choice to address this question. Organs are periodically initiated at the SAM at specific spatial positions and this spatiotemporal pattern defines phyllotaxis. Accumulation of the plant hormone auxin triggers organ initiation, whereas auxin depletion around organs generates inhibitory fields that are thought to be sufficient to maintain these patterns and their dynamics. Here we show that another type of hormone-based inhibitory fields, generated directly downstream of auxin by intercellular movement of the cytokinin signalling inhibitor ARABIDOPSIS HISTIDINE PHOSPHOTRANSFER PROTEIN 6 (AHP6), is involved in regulating phyllotactic patterns. We demonstrate that AHP6-based fields establish patterns of cytokinin signalling in the meristem that contribute to the robustness of phyllotaxis by imposing a temporal sequence on organ initiation. Our findings indicate that not one but two distinct hormone-based fields may be required for achieving temporal precision during formation of reiterative structures at the SAM, thus indicating an original mechanism for providing robustness to a dynamic developmental system.

A New Phenotyping Pipeline Reveals Three Types of Lateral Roots and a Random Branching Pattern in Two Cereals.

Recent progress in root phenotyping has focused mainly on increasing throughput for genetic studies, while identifying root developmental patterns has been comparatively underexplored. We introduce a new phenotyping pipeline for producing high-quality spatiotemporal root system development data and identifying developmental patterns within these data. The SmartRoot image-analysis system and temporal and spatial statistical models were applied to two cereals, pearl millet (Pennisetum glaucum) and maize (Zea mays). Semi-Markov switching linear models were used to cluster lateral roots based on their growth rate profiles. These models revealed three types of lateral roots with similar characteristics in both species. The first type corresponds to fast and accelerating roots, the second to rapidly arrested roots, and the third to an intermediate type where roots cease elongation after a few days. These types of lateral roots were retrieved in different proportions in a maize mutant affected in auxin signaling, while the first most vigorous type was absent in maize plants exposed to severe shading. Moreover, the classification of growth rate profiles was mirrored by a ranking of anatomical traits in pearl millet. Potential dependencies in the succession of lateral root types along the primary root were then analyzed using variable-order Markov chains. The lateral root type was not influenced by the shootward neighbor root type or by the distance from this root. This random branching pattern of primary roots was remarkably conserved, despite the high variability of root systems in both species. Our phenotyping pipeline opens the door to exploring the genetic variability of lateral root developmental patterns.

Identifying phenological phases in strawberry using multiple change-point models.

Plant development studies often generate data in the form of multivariate time series, each variable corresponding to a count of newly emerged organs for a given development process. These phenological data often exhibit highly structured patterns, and the aim of this study was to identify such patterns in cultivated strawberry. Six strawberry genotypes were observed weekly for their course of emergence of flowers, leaves, and stolons during 7 months. We assumed that these phenological series take the form of successive phases, synchronous between individuals. We applied univariate multiple change-point models for the identification of flowering, vegetative development, and runnering phases, and multivariate multiple change-point models for the identification of consensus phases for these three development processes. We showed that the flowering and the runnering processes are the main determinants of the phenological pattern. On this basis, we propose a typology of the six genotypes in the form of a hierarchical classification. This study introduces a new longitudinal data modeling approach for the identification of phenological phases in plant development. The focus was on development variables but the approach can be directly extended to growth variables and to multivariate series combining growth and development variables.

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.

Are compound leaves more complex than simple ones? A multi-scale analysis.

Background and Aims: The question of which cellular mechanisms determine the variation in leaf size has been addressed mainly in plants with simple leaves. It is addressed here in tomato taking into consideration the expected complexity added by the several lateral appendages making up the compound leaf, the leaflets. Methods: Leaf and leaflet areas, epidermal cell number and areas, and endoreduplication (co-) variations were analysed in Solanum lycopersicum considering heteroblastic series in a wild type (Wva106) and an antisense mutant, the Pro35S:Slccs52AAS line, and upon drought treatments. All plants were grown in an automated phenotyping platform, PHENOPSIS, adapted to host plants grown in 7 L pots. Key Results: Leaf area, leaflet area and cell number increased with leaf rank until reaching a plateau. In contrast, cell area slightly decreased and endoreduplication did not follow any trend. In the transgenic line, leaf area, leaflet areas and cell number of basal leaves were lower than in the wild type, but higher in upper leaves. Reciprocally, cell area was higher in basal leaves and lower in upper leaves. When scaled up at the whole sympodial unit, all these traits did not differ significantly between the transgenic line and the wild type. In response to drought, leaf area was reduced, with a clear dose effect that was also reported for all size-related traits, including endoreduplication. Conclusions: These results provide evidence that all leaflets have the same cellular phenotypes as the leaf they belong to. Consistent with results reported for simple leaves, they show that cell number rather than cell size determines the final leaf areas and that endoreduplication can be uncoupled from leaf and cell sizes. Finally, they re-question a whole-plant control of cell division and expansion in leaves when the Wva106 and the Pro35S:Slccs52AAS lines are compared.

Integrative models for joint analysis of shoot growth and branching patterns.

Plants exhibit dependences between shoot growth and branching that generate highly structured patterns. The characterization of the patterning mechanism is still an open issue because of the developmental processes involved with both succession of events (e.g. internode elongation, axillary shoot initiation and elongation) and complex dependences among neighbouring positions along the parent shoot. Statistical models called semi-Markov switching partitioned conditional generalized linear models were built on the basis of apple and pear tree datasets. In these models, the semi-Markov chain represents both the succession and lengths of branching zones, whereas the partitioned conditional generalized linear models represent the influence of parent shoot growth variables on axillary productions within each branching zone. Parent shoot growth variables were shown to influence specific developmental events. On this basis, the growth and branching patterns of two apple tree (Malus domestica) cultivars, as well as of pear trees (Pyrus spinosa) between two successive growing cycles, were compared. The proposed integrative statistical models were able to decipher the roles of successive developmental events in the growth and branching patterning mechanisms. These models could incorporate other parent shoot explanatory variables, such as the local curvature or the maximum growth rate of the leaf.

Identifying developmental phases in the Arabidopsis thaliana rosette using integrative segmentation models.

The change in leaf size and shape during ontogeny associated with heteroblastic development is a composite trait for which extensive spatiotemporal data can be acquired using phenotyping platforms. However, only part of the information contained in such data is exploited, and developmental phases are usually defined using a selected organ trait. We here introduce new methods for identifying developmental phases in the Arabidopsis rosette using various traits and minimum a priori assumptions. A pipeline of analysis was developed combining image analysis and statistical models to integrate morphological, shape, dimensional and expansion dynamics traits for the successive leaves of the Arabidopsis rosette. Dedicated segmentation models called semi-Markov switching models were built for selected genotypes in order to identify rosette developmental phases. Four successive developmental phases referred to as seedling, juvenile, transition and adult were identified for the different genotypes. We show that the degree of covering of the leaf abaxial surface with trichomes is insufficient to define these developmental phases. Using our pipeline of analysis, we were able to identify the supplementary seedling phase and to uncover the structuring role of various leaf traits. This enabled us to compare on a more objective basis the vegetative development of Arabidopsis mutants.

Identification of successive flowering phases highlights a new genetic control of the flowering pattern in strawberry.

The genetic control of the switch between seasonal and perpetual flowering has been deciphered in various perennial species. However, little is known about the genetic control of the dynamics of perpetual flowering, which changes abruptly at well-defined time instants during the growing season. Here, we characterize the perpetual flowering pattern and identify new genetic controls of this pattern in the cultivated strawberry. Twenty-one perpetual flowering strawberry genotypes were phenotyped at the macroscopic scale for their course of emergence of inflorescences and stolons during the growing season. A longitudinal analysis based on the segmentation of flowering rate profiles using multiple change-point models was conducted. The flowering pattern of perpetual flowering genotypes takes the form of three or four successive phases: an autumn-initiated flowering phase, a flowering pause, and a single stationary perpetual flowering phase or two perpetual flowering phases, the second one being more intense. The genetic control of flowering was analysed by quantitative trait locus mapping of flowering traits based on these flowering phases. We showed that the occurrence of a fourth phase of intense flowering is controlled by a newly identified locus, different from the locus FaPFRU, controlling the switch between seasonal and perpetual flowering behaviour. The role of this locus was validated by the analysis of data obtained previously during six consecutive years.

Analysing growth and development of plants jointly using developmental growth stages.

BACKGROUND AND AIMS: Plant growth, the increase of organ dimensions over time, and development, the change in plant structure, are often studied as two separate processes. However, there is structural and functional evidence that these two processes are strongly related. The aim of this study was to investigate the co-ordination between growth and development using mango trees, which have well-defined developmental stages. METHODS: Developmental stages, determined in an expert way, and organ sizes, determined from objective measurements, were collected during the vegetative growth and flowering phases of two cultivars of mango, Mangifera indica. For a given cultivar and growth unit type (either vegetative or flowering), a multistage model based on absolute growth rate sequences deduced from the measurements was first built, and then growth stages deduced from the model were compared with developmental stages. KEY RESULTS: Strong matches were obtained between growth stages and developmental stages, leading to a consistent definition of integrative developmental growth stages. The growth stages highlighted growth asynchronisms between two topologically connected organs, namely the vegetative axis and its leaves. CONCLUSIONS: Integrative developmental growth stages emphasize that developmental stages are closely related to organ growth rates. The results are discussed in terms of the possible physiological processes underlying these stages, including plant hydraulics, biomechanics and carbohydrate partitioning.

Differences in proleptic and epicormic shoot structures in relation to water deficit and growth rate in almond trees (Prunus dulcis).

BACKGROUND AND AIMS: Shoot characteristics differ depending on the meristem tissue that they originate from and environmental conditions during their development. This study focused on the effects of plant water status on axillary meristem fate and flowering patterns along proleptic and epicormic shoots, as well as on shoot growth rates on 'Nonpareil' almond trees (Prunus dulcis). The aims were (1) to characterize the structural differences between proleptic and epicormic shoots, (2) to determine whether water deficits modify shoot structures differently depending on shoot type, and (3) to determine whether shoot structures are related to shoot growth rates. METHODS: A hidden semi-Markov model of the axillary meristem fate and number of flower buds per node was built for two shoot types growing on trees exposed to three plant water status treatments. The models segmented observed shoots into successive homogeneous zones, which were compared between treatments. Shoot growth rates were calculated from shoot extension measurements made during the growing season. KEY RESULTS: Proleptic shoots had seven successive homogeneous zones while epicormic shoots had five zones. Shoot structures were associated with changes in growth rate over the season. Water deficit (1) affected the occurrence and lengths of the first zones of proleptic shoots, but only the occurrence of the third zone was reduced in epicormic shoots; (2) had a minor effect on zone flowering patterns and did not modify shoot or zone composition of axillary meristem fates; and (3) reduced growth rates, although patterns over the season were similar among treatments. CONCLUSIONS: Two meristem types, with different latency durations, produced shoots with different growth rates and distinct structures. Differences between shoot type structure responses to water deficit appeared to reflect their ontogenetic characteristics and/or resource availability for their development. Tree water deficit appeared to stimulate a more rapid progression through ontogenetic states.

Exploring bud dormancy completion with a combined architectural and phenological analysis: the case of apple trees in contrasting winter temperature conditions.

PREMISE OF THE STUDY: The branching pattern and phenology of trees result from interactions between the tree's genetic constitution and environmental conditions. Temperature strongly affects the duration of bud dormancy and further shoot growth. Our hypothesis was that shoot architecture is strongly affected by winter temperatures determining both the position and budburst of vegetative laterals with a lower effect on their outgrowth. METHODS: The study was conducted on four apple cultivars characterized by various chilling requirements and grown in two contrasting winter temperature conditions. A two-step approach was designed to quantify at the shoot scale first the branching pattern and second two phenological stages of vegetative laterals, budburst and outgrowth. A categorical variable, the branching zone, was built to summarize the lateral position along the shoot. It was integrated into the phenological analysis as a factor together with the cultivar and the winter temperature. KEY RESULTS: Temperature had a main effect on the distribution of vegetative laterals along the shoot. It also strongly affected budburst, which was also affected by the cultivar and the branching zone. The outgrowth of the lateral was not significantly affected by temperature but was significantly affected by the cultivar and the branching zone. Furthermore, the delayed senescence and subsequent leaf persistence during winter, characterizing the apple tree in the mild winter temperature condition, had only a weak effect on the distribution of vegetative laterals and on budburst and lateral outgrowth. CONCLUSIONS: The actual shoot architecture and budburst result from an ordered sequence of events with a pivotal role of winter temperatures on the dormancy completion of individual lateral buds. Endogenous factors related to the cultivar branching pattern overtake the temperature effect on the lateral outgrowth.

Glutaredoxin regulation of primary root growth is associated with early drought stress tolerance in pearl millet.

Seedling root traits impact plant establishment under challenging environments. Pearl millet is one of the most heat and drought tolerant cereal crops that provides a vital food source across the sub-Saharan Sahel region. Pearl millet's early root system features a single fast-growing primary root which we hypothesize is an adaptation to the Sahelian climate. Using crop modeling, we demonstrate that early drought stress is an important constraint in agrosystems in the Sahel where pearl millet was domesticated. Furthermore, we show that increased pearl millet primary root growth is correlated with increased early water stress tolerance in field conditions. Genetics including genome-wide association study and quantitative trait loci (QTL) approaches identify genomic regions controlling this key root trait. Combining gene expression data, re-sequencing and re-annotation of one of these genomic regions identified a glutaredoxin-encoding gene PgGRXC9 as the candidate stress resilience root growth regulator. Functional characterization of its closest Arabidopsis homolog AtROXY19 revealed a novel role for this glutaredoxin (GRX) gene clade in regulating cell elongation. In summary, our study suggests a conserved function for GRX genes in conferring root cell elongation and enhancing resilience of pearl millet to its Sahelian environment.

Pearl millet is a staple food for over 90 million people living in regions of Africa and India that typically experience high temperatures and little rainfall. It was domesticated about 4,500 years ago in the Sahel region of West Africa and is one of the most heat and drought tolerant cereal crops worldwide. In most plants, organs known as roots absorb water and essential nutrients from the soil. Young pearl millet plants develop a fast-growing primary root, but it is unclear how this unique feature helps the crop to grow in hot and dry conditions. Using weather data collected from the Sahel over a 20-year period, Fuente, Grondin et al. predicted by modelling that early drought stress is the major factor limiting pearl millet growth and yield in this region. Field experiments found that plants with primary roots that grow faster within soil were better at tolerating early drought than those with slower growing roots. Further work using genetic approaches revealed that a gene known as PgGRXC9 promotes the growth of the primary root. To better understand how this gene works, the team examined a very similar gene in a well-studied model plant known as Arabidopsis. This suggested that PgGRXC9 helps the primary root to grow by stimulating cell elongation within the root. Since it is well adapted to dry conditions, pearl millet is expected to play an important role in helping agriculture adjust to climate change. The findings of Fuente, Grondin et al. may be used by plant breeders to create more resilient and productive varieties of pearl millet.

Deciphering structural and temporal interplays during the architectural development of mango trees.

Plant architecture is commonly defined by the adjacency of organs within the structure and their properties. Few studies consider the effect of endogenous temporal factors, namely phenological factors, on the establishment of plant architecture. This study hypothesized that, in addition to the effect of environmental factors, the observed plant architecture results from both endogenous structural and temporal components, and their interplays. Mango tree, which is characterized by strong phenological asynchronisms within and between trees and by repeated vegetative and reproductive flushes during a growing cycle, was chosen as a plant model. During two consecutive growing cycles, this study described vegetative and reproductive development of 20 trees submitted to the same environmental conditions. Four mango cultivars were considered to assess possible cultivar-specific patterns. Integrative vegetative and reproductive development models incorporating generalized linear models as components were built. These models described the occurrence, intensity, and timing of vegetative and reproductive development at the growth unit scale. This study showed significant interplays between structural and temporal components of plant architectural development at two temporal scales. Within a growing cycle, earliness of bud burst was highly and positively related to earliness of vegetative development and flowering. Between growing cycles, flowering growth units delayed vegetative development compared to growth units that did not flower. These interplays explained how vegetative and reproductive phenological asynchronisms within and between trees were generated and maintained. It is suggested that causation networks involving structural and temporal components may give rise to contrasted tree architectures.

New insights for estimating the genetic value of segregating apple progenies for irregular bearing during the first years of tree production.

Because irregular bearing generates major agronomic issues in fruit-tree species, particularly in apple, the selection of regular cultivars is desirable. Here, we aimed to define methods and descriptors allowing a diagnostic for bearing behaviour during the first years of tree maturity, when tree production is increasing. Flowering occurrences were collected at whole-tree and (annual) shoot scales on a segregating apple population. At both scales, the number of inflorescences over the years was modelled. Two descriptors were derived from model residuals: a new biennial bearing index, based on deviation around yield trend over years and an autoregressive coefficient, which represents dependency between consecutive yields. At the shoot scale, entropy was also considered to represent the within-tree flowering synchronicity. Clusters of genotypes with similar bearing behaviours were built. Both descriptors at the whole-tree and shoot scales were consistent for most genotypes and were used to discriminate regular from biennial and irregular genotypes. Quantitative trait loci were detected for the new biennial bearing index at both scales. Combining descriptors at a local scale with entropy showed that regular bearing at the tree scale may result from different strategies of synchronization in flowering at the local scale. The proposed methods and indices open an avenue to quantify bearing behaviour during the first years of tree maturity and to capture genetic variations. Their extension to other progenies and species, possible variants of descriptors, and their use in breeding programmes considering a limited number of years or fruit yields are discussed.

Pattern identification and characterization reveal permutations of organs as a key genetically controlled property of post-meristematic phyllotaxis.

In vascular plants, the arrangement of organs around the stem generates geometric patterns called phyllotaxis. In the model plant, Arabidopsis thaliana, as in the majority of species, single organs are initiated successively at a divergence angle from the previous organ close to the canonical angle of 137.5°, producing a Fibonacci spiral. Given that little is known about the robustness of these geometric arrangements, we undertook to characterize phyllotaxis by measuring divergence angles between organs along the stems of wild-type and specific mutant plants with obvious defects in phyllotaxis. Sequences of measured divergence angles exhibit segments of non-canonical angles in both genotypes, albeit to a far greater extent in the mutant. We thus designed a pipeline of methods for analyzing these perturbations. The latent structure models used in this pipeline combine a non-observable model representing perturbation patterns (either a variable-order Markov chain or a combinatorial model) with von Mises distributions representing divergence angle uncertainty. We show that the segments of non-canonical angles in both wild-type and mutant plants can be explained by permutations in the order of insertion along the stem of two or three consecutive organs. The number of successive organs between two permutations reveals specific patterns that depend on the nature of the preceding permutation (2- or 3-permutation). We also highlight significant individual deviations from 137.5° in the level of baseline segments and a marked relationship between permutation of organs and defects in the elongation of the internodes between these organs. These results demonstrate that permutations are an intrinsic property of spiral phyllotaxis and that their occurrence is genetically regulated.

The auxin signalling network translates dynamic input into robust patterning at the shoot apex.

The plant hormone auxin is thought to provide positional information for patterning during development. It is still unclear, however, precisely how auxin is distributed across tissues and how the hormone is sensed in space and time. The control of gene expression in response to auxin involves a complex network of over 50 potentially interacting transcriptional activators and repressors, the auxin response factors (ARFs) and Aux/IAAs. Here, we perform a large-scale analysis of the Aux/IAA-ARF pathway in the shoot apex of Arabidopsis, where dynamic auxin-based patterning controls organogenesis. A comprehensive expression map and full interactome uncovered an unexpectedly simple distribution and structure of this pathway in the shoot apex. A mathematical model of the Aux/IAA-ARF network predicted a strong buffering capacity along with spatial differences in auxin sensitivity. We then tested and confirmed these predictions using a novel auxin signalling sensor that reports input into the signalling pathway, in conjunction with the published DR5 transcriptional output reporter. Our results provide evidence that the auxin signalling network is essential to create robust patterns at the shoot apex.

Identifying Developmental Patterns in Structured Plant Phenotyping Data.

Technological breakthroughs concerning both sensors and robotized plant phenotyping platforms have totally renewed the plant phenotyping paradigm in the last two decades. This has impacted both the nature and the throughput of data with the availability of data at high-throughput from the tissular to the whole plant scale. Sensor outputs often take the form of 2D or 3D images or time series of such images from which traits are extracted while organ shapes, shoot or root system architectures can be deduced. Despite this change of paradigm, many phenotyping studies often ignore the structure of the plant and therefore loose the information conveyed by the temporal and spatial patterns emerging from this structure. The developmental patterns of plants often take the form of succession of well-differentiated phases, stages or zones depending on the temporal, spatial or topological indexing of data. This entails the use of hierarchical statistical models for their identification.The objective here is to show potential approaches for analyzing structured plant phenotyping data using state-of-the-art methods combining probabilistic modeling, statistical inference and pattern recognition. This approach is illustrated using five different examples at various scales that combine temporal and topological index parameters, and development and growth variables obtained using prospective or retrospective measurements.

Deciphering the developmental plasticity of walnut saplings in relation to climatic factors and light environment.

Developmental plasticity, the acclimation of plants to their local environment, is known to be crucial for the fitness of perennial organisms such as trees. However, deciphering the many possible developmental and environmental influences involved in such plasticity in natural conditions requires dedicated statistical models integrating developmental phases, environmental factors, and interindividual heterogeneity. These models should be able to analyse retrospective data (number of leaves or length of annual shoots along the main stem in the present case). In this study Markov switching linear mixed models were applied to the analysis of the developmental plasticity of walnut saplings during the establishment phase in a mixed Mediterranean forest. In the Markov switching linear mixed models estimated from walnut data sets, the underlying Markov chain represents both the succession and lengths of growth phases, while the linear mixed models represent both the influence of climatic factors and interindividual heterogeneity within each growth phase. On the basis of these integrative statistical models, it is shown that walnut saplings have an opportunistic mode of development that is primarily driven by the changing light environment. In particular, light availability explains the ability of a tree to reach a phase of strong growth where the first branches can appear. It is also shown that growth fluctuation amplitudes in response to climatic factors increased while interindividual heterogeneity decreased during tree development.