Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs.

The effects of plants on the biosphere, atmosphere and geosphere are key determinants of terrestrial ecosystem functioning. However, despite substantial progress made regarding plant belowground components, we are still only beginning to explore the complex relationships between root traits and functions. Drawing on the literature in plant physiology, ecophysiology, ecology, agronomy and soil science, we reviewed 24 aspects of plant and ecosystem functioning and their relationships with a number of root system traits, including aspects of architecture, physiology, morphology, anatomy, chemistry, biomechanics and biotic interactions. Based on this assessment, we critically evaluated the current strengths and gaps in our knowledge, and identify future research challenges in the field of root ecology. Most importantly, we found that belowground traits with the broadest importance in plant and ecosystem functioning are not those most commonly measured. Also, the estimation of trait relative importance for functioning requires us to consider a more comprehensive range of functionally relevant traits from a diverse range of species, across environments and over time series. We also advocate that establishing causal hierarchical links among root traits will provide a hypothesis-based framework to identify the most parsimonious sets of traits with the strongest links on functions, and to link genotypes to plant and ecosystem functioning.

A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements.

In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I-VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers' views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning.

Modelling time variations of root diameter and elongation rate as related to assimilate supply and demand.

In a given root system, individual roots usually exhibit a rather homogeneous tip structure although highly different diameters and growth patterns, and this diversity is of prime importance in the definition of the whole root system architecture and foraging characteristics. In order to represent and predict this diversity, we built a simple and generic model at root tip level combining structural and functional knowledge on root elongation. The tip diameter, reflecting meristem size, is used as a driving variable of elongation. It varies, in response to the fluctuations of photo-assimilate availability, between two limits (minimal and maximal diameter). The elongation rate is assumed to be dependent on the transient value of the diameter. Elongation stops when the tip reaches the minimal diameter. The model could satisfactorily reproduce patterns of root elongation and tip diameter changes observed in various species at different scales. Although continuous, the model could generate divergent root classes as classically observed within populations of lateral roots. This model should help interpret the large plasticity of root elongation patterns which can be obtained in response to different combinations of endogenous and exogenous factors. The parameters could be used in phenotyping the root system.

Simulating the effects of water limitation on plant biomass using a 3D functional-structural plant model of shoot and root driven by soil hydraulics.

BACKGROUND AND AIMS: Improved modelling of carbon assimilation and plant growth to low soil moisture requires evaluation of underlying mechanisms in the soil, roots, and shoots. The feedback between plants and their local environment throughout the whole spectrum soil-root-shoot-environment is crucial to accurately describe and evaluate the impact of environmental changes on plant development. This study presents a 3D functional structural plant model, in which shoot and root growth are driven by radiative transfer, photosynthesis, and soil hydrodynamics through different parameterisation schemes relating soil water deficit and carbon assimilation. The new coupled model is used to evaluate the impact of soil moisture availability on plant productivity for two different groups of flowering plants under different spatial configurations. METHODS: In order to address different aspects of plant development due to limited soil water availability, a 3D FSP model including root, shoot, and soil was constructed by linking three different well-stablished models of airborne plant, root architecture, and reactive transport in the soil. Different parameterisation schemes were used in order to integrate photosynthetic rate with root water uptake within the coupled model. The behaviour of the model was assessed on how the growth of two different types of plants, i.e. monocot and dicot, is impacted by soil water deficit under different competitive conditions: isolated (no competition), intra, and interspecific competition. KEY RESULTS: The model proved to be capable of simulating carbon assimilation and plant development under different growing settings including isolated monocots and dicots, intra, and interspecific competition. The model predicted that (1) soil water availability has a larger impact on photosynthesis than on carbon allocation; (2) soil water deficit has an impact on root and shoot biomass production by up to 90 % for monocots and 50 % for dicots; and (3) the improved dicot biomass production in interspecific competition was highly related to root depth and plant transpiration. CONCLUSIONS: An integrated model of 3D shoot architecture and biomass development with a 3D root system representation, including light limitation and water uptake considering soil hydraulics, was presented. Plant-plant competition and regulation on stomatal conductance to drought were able to be predicted by the model. In the cases evaluated here, water limitation impacted plant growth almost 10 times more than the light environment.

Connecting the dots between computational tools to analyse soil-root water relations.

In recent years, many computational tools, such as image analysis, data management, process-based simulation, and upscaling tools, have been developed to help quantify and understand water flow in the soil-root system, at multiple scales (tissue, organ, plant, and population). Several of these tools work together or at least are compatible. However, for the uninformed researcher, they might seem disconnected, forming an unclear and disorganized succession of tools. In this article, we show how different studies can be further developed by connecting them to analyse soil-root water relations in a comprehensive and structured network. This 'explicit network of soil-root computational tools' informs readers about existing tools and helps them understand how their data (past and future) might fit within the network. We also demonstrate the novel possibilities of scale-consistent parameterizations made possible by the network with a set of case studies from the literature. Finally, we discuss existing gaps in the network and how we can move forward to fill them.

Seeking stable traits to characterize the root system architecture. Study on 60 species located at two sites in natura.

Background and Aims: In several disciplines, identifying relevant root traits to characterize the root system architecture of species or genotypes is a crucial step. To address this question, we analysed the inter-specific variations of root architectural traits in two contrasting environments. Methods: We sampled 60 species in natura, at two sites, each presenting homogeneous soil conditions. We estimated for each species and site a set of five traits used for the modelling of the root system architecture: extreme tip diameters (Dmin and Dmax), relative diameter range (Drange), mean inter-branch distance (IBD) and dominance slope between the diameters of parent and lateral roots (DlDm). Key Results: The five traits presented a highly significant species effect, explaining between 77 and 98 % of the total variation. Dmin, Dmax and Drange were particularly determined by the species, while DlDm and IBD exhibited a higher percentage of environmental variations. These traits make it possible to confirm two main axes of variation: 'fineness-density' (defined by Dmin and IBD) and 'dominance-heterorhizy' (DlDm and Drange), that together accounted for 84 % of the variations observed. Conclusions: We confirmed the interest of these traits in the characterization of the root system architecture in ecology and genetics, and suggest using them to enrich the 'root economic spectrum'.

Branching patterns of root systems: comparison of monocotyledonous and dicotyledonous species.

BACKGROUND AND AIMS: Acropetal root branching is a major process which increases the number of growing tips and distributes their growth potential within the whole root system. METHODS: Using a method presented in a recent paper, the defined branching traits were estimated in 140 different species, and the branching patterns of monocots (45 species) and dicots (95 species) were compared. KEY RESULTS: It was checked that the method also applied to monocots (not considered in the previous paper), and that all traits could be estimated in each species. Variations of most traits were even larger for monocots than for dicots. Systematic differences appeared between these two groups: monocots tended to have a larger range in apical diameters (stronger heterorhizy), with both finer and thicker roots; the diameters of their lateral roots were also more variable; their roots exerted a stronger dominance over lateral branches. Altogether, species exhibited two main dependencies among their traits that were illustrated using two axes: (1) the 'fineness-density' axis separated the species which develop very fine roots and branch densely, from species without fine roots which space out their branches; and (2) the 'dominance-heterorhizy' axis separated the species according to the range in their apical diameter which was positively correlated to the level of dominance of mother roots over their branches. Both axes and correlations were remarkably similar for monocots and dicots. CONCLUSIONS: Beyond the overall typology, this study went on to validate the phenotyping method in Natura, and showed its potential to characterize the differences in groups of species.

Relationships between root diameter, root length and root branching along lateral roots in adult, field-grown maize.

BACKGROUND AND AIMS: Root diameter, especially apical diameter, plays an important role in root development and function. The variation in diameter between roots, and along roots, affects root structure and thus the root system's overall foraging performance. However, the effect of diameter variation on root elongation, branching and topological connections has not been examined systematically in a population of high-order roots, nor along the roots, especially for mature plants grown in the field. METHODS: A method combining both excavation and analysis was applied to extract and quantify root architectural traits of adult, field-grown maize plants. The relationships between root diameter and other root architectural characteristics are analysed for two maize cultivars. KEY RESULTS: The basal diameter of the lateral roots (orders 1-3) was highly variable. Basal diameter was partly determined by the diameter of the bearing segment. Basal diameter defined a potential root length, but the lengths of most roots fell far short of this. This was explained partly by differences in the pattern of diameter change along roots. Diameter tended to decrease along most roots, with the steepness of the gradient of decrease depending on basal diameter. The longest roots were those that maintained (or sometimes increased) their diameters during elongation. The branching density (cm(-1)) of laterals was also determined by the diameter of the bearing segment. However, the location of this bearing segment along the mother root was also involved - intermediate positions were associated with higher densities of laterals. CONCLUSIONS: The method used here allows us to obtain very detailed records of the geometry and topology of a complex root system. Basal diameter and the pattern of diameter change along a root were associated with its final length. These relationships are especially useful in simulations of root elongation and branching in source-sink models.

Modelling the root system architecture of Poaceae. Can we simulate integrated traits from morphological parameters of growth and branching?

Our objective was to calibrate a model of the root system architecture on several Poaceae species and to assess its value to simulate several 'integrated' traits measured at the root system level: specific root length (SRL), maximum root depth and root mass. We used the model ArchiSimple, made up of sub-models that represent and combine the basic developmental processes, and an experiment on 13 perennial grassland Poaceae species grown in 1.5-m-deep containers and sampled at two different dates after planting (80 and 120 d). Model parameters were estimated almost independently using small samples of the root systems taken at both dates. The relationships obtained for calibration validated the sub-models, and showed species effects on the parameter values. The simulations of integrated traits were relatively correct for SRL and were good for root depth and root mass at the two dates. We obtained some systematic discrepancies that were related to the slight decline of root growth in the last period of the experiment. Because the model allowed correct predictions on a large set of Poaceae species without global fitting, we consider that it is a suitable tool for linking root traits at different organisation levels.

Branching patterns of root systems: quantitative analysis of the diversity among dicotyledonous species.

BACKGROUND AND AIMS: Root branching, and in particular acropetal branching, is a common and important developmental process for increasing the number of growing tips and defining the distribution of their meristem size. This study presents a new method for characterizing the results of this process in natura from scanned images of young, branched parts of excavated roots. The method involves the direct measurement or calculation of seven different traits. METHODS: Young plants of 45 species of dicots were sampled from fields and gardens with uniform soils. Roots were separated, scanned and then measured using ImageJ software to determine seven traits related to root diameter and interbranch distance. RESULTS: The traits exhibited large interspecific variations, and covariations reflecting trade-offs. For example, at the interspecies level, the spacing of lateral roots (interbranch distance along the parent root) was strongly correlated to the diameter of the finest roots found in the species, and showed a continuum between two opposite strategies: making dense and fine lateral roots, or thick and well-spaced laterals. CONCLUSIONS: A simple method is presented for classification of branching patterns in roots that allows relatively quick sampling and measurements to be undertaken. The feasibilty of the method is demonstrated for dicotyledonous species and it has the potential to be developed more broadly for other species and a wider range of enivironmental conditions.

[Onco'sport testing by patients: Lived experience of patients with cancer practicing adapted physical activity as part of the Onco'sport program, a qualitative study]. / Onco'sport à l'épreuve des patients : expérience vécue des patients atteints d'un cancer pratiquant de l'activité physique adaptée dans le cadre du programme Onco'sport, une étude qualitative.

INTRODUCTION: Physical activity is a major determinant in the prevention of chronic diseases, equally on the side effects of treatments and the consequences of the disease. It improves quality of life, but also reduces morbidity and mortality, and therefore health expenses. A sedentary lifestyle is the fourth cause of premature death in the world, in the context of chronic non-communicable diseases. In France, the prescription for adapted physical activity (APA) has been included in the law since 2016. With the development of "Maisons Sport santé", the Onco'sport program was developed to enable people affected by cancer to participate in adapted physical activity. The objective of our work is to explore the lived experience of cancer patients practicing adapted physical activity as part of the Onco'sport program. METHODS: This is a qualitative study of 10 semi-directed individual interviews with patients participating in the Onco'sport program, recruited from the "Maison Sport Santé" from Nîmes and the association "Les Roses du Gard". A phenomenological analysis was carried out with a semiopragmatic approach. RESULTS: For all participants, the APA through a program provides professional supervision of Physical Activity, influences adherence and builds confidence. This program is at the origin of changes in lifestyle habits and improves the relationship with illness and their cancer thanks to the physical and psychological benefits felt. Thus, APA appears as a full-fledged supportive care which requires informing patients and promoting it so that access is facilitated and becomes a standard. Health professionals including general practitioners do not currently have an important place in the promotion of APA, and patients most often obtain documentation on their own or through associations of patient. CONCLUSION: An APA program like Onco'sport is often the cause of a return to physical activity in patients, brings overall well-being and changes lifestyle habits. It seems important to promote physical activity to patients but also to the general population, given the benefits. This promotion involves easier access to this type of supervised program, financial support and better training of health professionals.

A novel image-analysis toolbox enabling quantitative analysis of root system architecture.

We present in this paper a novel, semiautomated image-analysis software to streamline the quantitative analysis of root growth and architecture of complex root systems. The software combines a vectorial representation of root objects with a powerful tracing algorithm that accommodates a wide range of image sources and quality. The root system is treated as a collection of roots (possibly connected) that are individually represented as parsimonious sets of connected segments. Pixel coordinates and gray level are therefore turned into intuitive biological attributes such as segment diameter and orientation as well as distance to any other segment or topological position. As a consequence, user interaction and data analysis directly operate on biological entities (roots) and are not hampered by the spatially discrete, pixel-based nature of the original image. The software supports a sampling-based analysis of root system images, in which detailed information is collected on a limited number of roots selected by the user according to specific research requirements. The use of the software is illustrated with a time-lapse analysis of cluster root formation in lupin (Lupinus albus) and an architectural analysis of the maize (Zea mays) root system. The software, SmartRoot, is an operating system-independent freeware based on ImageJ and relies on cross-platform standards for communication with data-analysis software.

Links between root developmental traits and foraging performance.

We designed a simple dynamic and stochastic architectural model with six parameters to link the foraging performance of root systems to their developmental processes. Soil foraging was quantified by the volume enveloping the roots until a given uptake distance. Many simulated architectures were obtained by combining four different values for each parameter. The rate of soil colonization was mainly defined by individual root elongation rates and interbranch distances. Less intuitively, we showed that differentiation of elongation rates among the roots increased this colonization rate. Uptake efficiency--the ratio of the actual colonized volume to the volume of a unique cylinder with the same length and a radius corresponding to the uptake distance--declined with root system size. Nevertheless, large variations in efficiency existed among root systems for a given size, typically in a 4- to 10-fold range. Therefore, the 'efficiency gain' was defined as the deviation from the average trend in efficiency versus size. Between-root differentiation in elongation rates increased this gain. The level of hierarchy between mother and lateral roots, as well as the variation of elongation rates among lateral roots, was also shown to contribute to this optimization. Several parameter combinations could lead to similar efficiency gains.

Correlated responses of root growth and sugar concentrations to various defoliation treatments and rhythmic shoot growth in oak tree seedlings (Quercus pubescens).

BACKGROUND AND AIMS: To understand whether root responses to aerial rhythmic growth and contrasted defoliation treatments can be interpreted under the common frame of carbohydrate availability; root growth was studied in parallel with carbohydrate concentrations in different parts of the root system on oak tree seedlings. METHODS: Quercus pubescens seedlings were submitted to selective defoliation (removal of mature leaves, cotyledons or young developing leaves) at appearance of the second flush and collected 1, 5 or 10 d later for morphological and biochemical measurements. Soluble sugar and starch concentrations were measured in cotyledons and apical and basal root parts. KEY RESULTS: Soluble sugar concentration in the root apices diminished during the expansion of the second aerial flush and increased after the end of aerial growth in control seedlings. Starch concentration in cotyledons regularly decreased. Continuous removal of young leaves did not alter either root growth or apical sugar concentration. Starch storage in basal root segments was increased. After removal of mature leaves (and cotyledons), root growth strongly decreased. Soluble sugar concentration in the root apices drastically decreased and starch reserves in the root basal segments were emptied 5 d after defoliation, illustrating a considerable shortage in carbohydrates. Soluble sugar concentrations recovered 10 d after defoliation, after the end of aerial growth, suggesting a recirculation of sugar. No supplementary recourse to starch in cotyledons was observed. CONCLUSIONS: The parallel between apical sugar concentration and root growth patterns, and the correlations between hexose concentration in root apices and their growth rate, support the hypothesis that the response of root growth to aerial periodic growth and defoliation treatments is largely controlled by carbohydrate availability.

Genotypic diversity and plasticity of root system architecture to nitrogen availability in oilseed rape.

In the emerging new agricultural context, a drastic reduction in fertilizer usage is required. A promising way to maintain high crop yields while reducing fertilizer inputs is to breed new varieties with optimized root system architecture (RSA), designed to reach soil resources more efficiently. This relies on identifying key traits that underlie genotypic variability and plasticity of RSA in response to nutrient availability. The aim of our study was to characterize the RSA plasticity in response to nitrogen limitation of a set of contrasted oilseed rape genotypes, by using the ArchiSimple model parameters as screening traits. Eight accessions of Brassica napus were grown in long tubes in the greenhouse, under two contrasting levels of nitrogen availability. After plant excavation, roots were scanned at high resolution. Six RSA traits relative to root diameter, elongation rate and branching were measured, as well as nine growth and biomass allocation traits. The plasticity of each trait to nitrogen availability was estimated. Nitrogen-limited plants were characterized by a strong reduction in total biomass and leaf area. Even if the architecture traits were shown to be less plastic than allocation traits, significant nitrogen and genotype effects were highlighted on each RSA trait, except the root minimal diameter. Thus, the RSA of nitrogen-limited plants was primarily characterised by a reduced lateral root density, a smaller primary root diameter, associated with a stronger root dominance. Among the RSA traits measured, the inter-branch distance showed the highest plasticity with a level of 70%, in the same range as the most plastic allocation traits. This work suggests that lateral root density plays the key role in the adaptation of the root system to nitrogen availability and highlights inter-branch distance as a major target trait for breeding new varieties, better adapted to low input systems.

Genetic Variation in Root Architectural Traits in Lactuca and Their Roles in Increasing Phosphorus-Use-Efficiency in Response to Low Phosphorus Availability.

Low phosphorus (P) bioavailability in the soil and concerns over global P reserves have emphasized the need to cultivate plants that acquire and use P efficiently. Root architecture adaptation to low P can be variable depending on species or even genotypes. To assess the genetic variability of root architectural traits and their responses to low P in the Lactuca genus, we examined fourteen genotypes including wild species, ancient and commercial lettuce cultivars at low (LP, 0.1 mmol. L-1) and high P (HP, 1 mmol. L-1). Plants were grown in cylindrical pots adapted for the excavation and observation of root systems, with an inert substrate. We identified substantial genetic variation in all the investigated root traits, as well as an effect of P availability on these traits, except on the diameter of thinner roots. At low P, the main responses were a decrease in taproot diameter, an increase in taproot dominance over its laterals and an increase in the inter-branch distance. Although the genotype x P treatment effect was limited to root depth, we identified a tradeoff between the capacity to maintain a thick taproot at low P and the dominance of the taproot over its laterals. Regardless of the P level, the phosphorus-use-efficiency (PUE) varied among lettuce genotypes and was significantly correlated with total root biomass regardless of the P level. As taproot depth and maximum apical diameter were the principal determinants of total root biomass, the relative increase in PUE at low P was observed in genotypes that showed the thickest apical diameters and/or those whose maximal apical diameter was not severely decreased at low P availability. This pre-eminence of the taproot in the adaptation of Lactuca genotypes to low P contrasts with other species which rely more on lateral roots to adapt to P stress.

To what extent may changes in the root system architecture of Arabidopsis thaliana grown under contrasted homogenous nitrogen regimes be explained by changes in carbon supply? A modelling approach.

Root system architecture adapts to low nitrogen (N) nutrition. Some adaptations may be mediated by modifications of carbon (C) fluxes. The objective of this study was to test the hypothesis that changes in root system architecture under different N regimes may be accounted for by using simple hypotheses of C allocation within the root system of Arabidopsis thaliana. With that purpose, a model during vegetative growth was developed that predicted the main traits of root system architecture (total root length, lateral root number, and specific root length). Different experimental data sets crossing three C levels and two N homogenous nutrition levels were generated. Parameters were estimated from an experiment carried out under medium C and high N conditions. They were then checked under other CxN conditions. It was found that the model was able to simulate correctly C effects on root architecture in both high and low N nutrition conditions, with the same parameter values. It was concluded that C flux modifications explained the major part of root system adaptation to N supply, even if they were not sufficient to simulate some changes, such as specific root length.

Diel changes in nitrogen and carbon resource status and use for growth in young plants of tomato (Solanum lycopersicum).

BACKGROUND AND AIMS: Modellers often define growth as the development of plant structures from endogenous resources, thus making a distinction between structural (W(S)) and total (W) dry biomass, the latter being the sum of W(S) and the weight of storage compounds. In this study, short-term C and N reserves were characterized experimentally (forms, organ distribution, time changes) in relation to light and nutrition signals, and organ structural growth in response to reserve levels was evaluated. METHODS: Tomato plants (Solanum lycopersicum) were grown hydroponically in a growth room with a 12-h photoperiod and an adequate supply of NO(3)(-) (3 mol m(-3)). Three experiments were carried out 18 d after sowing: [NO(3)(-)] was either maintained at 3 mol m(-3), changed to 0.02 mol m(-3) or to 0 mol m(-3). Plants were sampled periodically throughout the light/dark cycles over 24-48 h. Organ W(S) was calculated from W together with the amount of different compounds that act as C and N resources, i.e. non-structural carbohydrates and carboxylates, nitrate and free amino acids. KEY RESULTS: With adequate nutrition, carbohydrates accumulated in leaves during light periods, when photosynthesis exceeded growth needs, but decreased at night when these sugars are the main source of C for growth. At the end of the night, carbohydrates were still high enough to fuel full-rate growth, as W(S) increased at a near constant rate throughout the light/dark cycle. When nitrate levels were restricted, C reserves increased, but [NO(3)(-)] decreased progressively in stems, which contain most of the plant N reserves, and rapidly in leaves and roots. This resulted in a rapid restriction of structural growth. CONCLUSIONS: Periodic darkness did not restrict growth because sufficient carbohydrate reserves accumulated during the light period. Structural growth, however, was very responsive to NO(3)(-) nutrition, because N reserves were mostly located in stems, which have limited nitrate reduction capacity.

Root system architecture: opportunities and constraints for genetic improvement of crops.

Abiotic stresses increasingly curtail crop yield as a result of global climate change and scarcity of water and nutrients. One way to minimize the negative impact of these factors on yield is to manipulate root system architecture (RSA) towards a distribution of roots in the soil that optimizes water and nutrient uptake. It is now established that most of the genetic variation for RSA is driven by a suite of quantitative trait loci. As we discuss here, marker-assisted selection and quantitative trait loci cloning for RSA are underway, exploiting genomic resources, candidate genes and the knowledge gained from Arabidopsis, rice and other crops. Nonetheless, efficient and accurate phenotyping, modelling and collaboration with breeders remain important challenges, particularly when defining ideal RSA for different crops and target environments.