Functional trade-offs are driven by coordinated changes among cell types in the wood of angiosperm trees from different climates.

Wood performs several functions to ensure tree survival and carbon allocation to a finite stem volume leads to trade-offs among cell types. It is not known to what extent these trade-offs modify functional trade-offs and if they are consistent across climates and evolutionary lineages. Twelve wood traits were measured in stems and coarse roots across 60 adult angiosperm tree species from temperate, Mediterranean and tropical climates. Regardless of climate, clear trade-offs occurred among cellular fractions, but did not translate into specific functional trade-offs. Wood density was negatively related to hydraulic conductivity (Kth ) in stems and roots, but was not linked to nonstructural carbohydrates (NSC), implying a functional trade-off between mechanical integrity and transport but not with storage. NSC storage capacity was positively associated with Kth in stems and negatively in roots, reflecting a potential role for NSC in the maintenance of hydraulic integrity in stems but not in roots. Results of phylogenetic analyses suggest that evolutionary histories cannot explain covariations among traits. Trade-offs occur among cellular fractions, without necessarily modifying trade-offs in function. However, functional trade-offs are driven by coordinated changes among xylem cell types depending on the dominant role of each cell type in stems and roots.

Non-structural carbohydrates and morphological traits of leaves, stems and roots from tree species in different climates.

OBJECTIVES: Carbon fixed during photosynthesis is exported from leaves towards sink organs as non-structural carbohydrates (NSC), that are a key energy source for metabolic processes in trees. In xylem, NSC are mostly stored as soluble sugars and starch in radial and axial parenchyma. The multi-functional nature of xylem means that cells possess several functions, including water transport, storage and mechanical support. Little is known about how NSC impacts xylem multi-functionality, nor how NSC vary among species and climates. We collected leaves, stem and root xylem from tree species growing in three climates and estimated NSC in each organ. We also measured xylem traits linked to hydraulic and mechanical functioning. DATA DESCRIPTION: The paper describes functional traits in leaves, stems and roots, including NSC, carbon, nitrogen, specific leaf area, stem and root wood density and xylem traits. Data are provided for up to 90 angiosperm species from temperate, Mediterranean and tropical climates. These data are useful for understanding the trade-offs in resource allocation from a whole-plant perspective, and to better quantify xylem structure and function related to water transportation, mechanical support and storage. Data will also give researchers keys to understanding the ability of trees to adjust to a changing climate.

Parenchyma fractions drive the storage capacity of nonstructural carbohydrates across a broad range of tree species.

PREMISE: Nonstructural carbohydrates (NSCs) play a key role in tree performance and functioning and are stored in radial and axial parenchyma (RAP) cells. Whether this relationship is altered among species and climates or is linked to functional traits describing xylem structure (wood density) and tree stature is not known. METHODS: In a systematic review, we collated data for NSC content and the proportion of RAP in stems for 68 tree species. To examine the relationships of NSCs and RAP with climatic factors and other functional traits, we also collected climatic data at each tree's location, as well as wood density and maximum height. A phylogenetic tree was constructed to examine the influence of species' evolutionary relationships on the associations among NSCs, RAP, and functional traits. RESULTS: Across all 68 tree species, NSCs were positively correlated with RAP and mean annual temperature, but relationships were only weakly significant in temperate species and angiosperms. When separating RAP into radial parenchyma (RP) and axial parenchyma (AP), both NSCs and wood density were positively correlated with RP but not with AP. Wood in taller trees was less dense and had lower RAP than in shorter trees, but height was not related to NSCs. CONCLUSIONS: In trees, NSCs are stored mostly in the RP fraction, which has a larger surface area in warmer climates. Additionally, NSCs were only weakly linked to wood density and tree height. Our analysis of evolutionary relationships demonstrated that RAP fractions and NSC content were always closely related across all 68 tree species, suggesting that RAP can act as a reliable proxy for potential NSC storage capacity in tree stems.

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.

Shifts in soil and plant functional diversity along an altitudinal gradient in the French Alps.

OBJECTIVES: Altitude integrates changes in environmental conditions that determine shifts in vegetation, including temperature, precipitation, solar radiation and edaphogenetic processes. In turn, vegetation alters soil biophysical properties through litter input, root growth, microbial and macrofaunal interactions. The belowground traits of plant communities modify soil processes in different ways, but it is not known how root traits influence soil biota at the community level. We collected data to investigate how elevation affects belowground community traits and soil microbial and faunal communities. This dataset comprises data from a temperate climate in France and a twin study was performed in a tropical zone in Mexico. DATA DESCRIPTION: The paper describes soil physical and chemical properties, climatic variables, plant community composition and species abundance, plant community traits, soil microbial functional diversity and macrofaunal abundance and diversity. Data are provided for six elevations (1400-2400 m) ranging from montane forest to alpine prairie. We focused on soil biophysical properties beneath three dominant plant species that structure local vegetation. These data are useful for understanding how shifts in vegetation communities affect belowground processes, such as water infiltration, soil aggregation and carbon storage. Data will also help researchers understand how plant communities adjust to a changing climate/environment.

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.

Belowground impacts of alpine woody encroachment are determined by plant traits, local climate, and soil conditions.

Global climate and land use change are causing woody plant encroachment in arctic, alpine, and arid/semi-arid ecosystems around the world, yet our understanding of the belowground impacts of this phenomenon is limited. We conducted a globally distributed field study of 13 alpine sites across four continents undergoing woody plant encroachment and sampled soils from both woody encroached and nearby herbaceous plant community types. We found that woody plant encroachment influenced soil microbial richness and community composition across sites based on multiple factors including woody plant traits, site level climate, and abiotic soil conditions. In particular, root symbiont type was a key determinant of belowground effects, as Nitrogen-fixing woody plants had higher soil fungal richness, while Ecto/Ericoid mycorrhizal species had higher soil bacterial richness and symbiont types had distinct soil microbial community composition. Woody plant leaf traits indirectly influenced soil microbes through their impact on soil abiotic conditions, primarily soil pH and C:N ratios. Finally, site-level climate affected the overall magnitude and direction of woody plant influence, as soil fungal and bacterial richness were either higher or lower in woody encroached versus herbaceous soils depending on mean annual temperature and precipitation. All together, these results document global impacts of woody plant encroachment on soil microbial communities, but highlight that multiple biotic and abiotic pathways must be considered to scale up globally from site- and species-level patterns. Considering both the aboveground and belowground effects of woody encroachment will be critical to predict future changes in alpine ecosystem structure and function and subsequent feedbacks to the global climate system.

Mechanical traits of fine roots as a function of topology and anatomy.

Background and Aims: Root mechanical traits, including tensile strength (Tr), tensile strain (εr) and modulus of elasticity (Er), are key functional traits that help characterize plant anchorage and the physical contribution of vegetation to landslides and erosion. The variability in these traits is high among tree fine roots and is poorly understood. Here, we explore the variation in root mechanical traits as well as their underlying links with morphological (diameter), architectural (topological order) and anatomical (stele and cortex sizes) traits. Methods: We investigated the four tropical tree species Pometia tomentosa, Barringtonia fusicarpa, Baccaurea ramiflora and Pittosporopsis kerrii in Xishuangbanna, Yunnan, China. For each species, we excavated intact, fresh, fine roots and measured mechanical and anatomical traits for each branching order. Key Results: Mechanical traits varied enormously among the four species within a narrow range of diameters (<2 mm): <0.1-65 MPa for Tr, 4-1135 MPa for Er and 0.4-37 % for εr. Across species, Tr and Er were strongly correlated with stele area ratio, which was also better correlated with topological order than with root diameter, especially at interspecific levels. Conclusions: Root topological order plays an important role in explaining variability in fine-root mechanical traits due to its reflection of root tissue development. Accounting for topological order when measuring fine-root traits therefore leads to greater empirical understanding of plant functions (e.g. anchorage) within and across species.

An evaluation of inexpensive methods for root image acquisition when using rhizotrons.

BACKGROUND: Belowground processes play an essential role in ecosystem nutrient cycling and the global carbon budget cycle. Quantifying fine root growth is crucial to the understanding of ecosystem structure and function and in predicting how ecosystems respond to climate variability. A better understanding of root system growth is necessary, but choosing the best method of observation is complex, especially in the natural soil environment. Here, we compare five methods of root image acquisition using inexpensive technology that is currently available on the market: flatbed scanner, handheld scanner, manual tracing, a smartphone application scanner and a time-lapse camera. Using the five methods, root elongation rate (RER) was measured for three months, on roots of hybrid walnut (Juglans nigra × Juglans regia L.) in rhizotrons installed in agroforests. RESULTS: When all methods were compared together, there were no significant differences in relative cumulative root length. However, the time-lapse camera and the manual tracing method significantly overestimated the relative mean diameter of roots compared to the three scanning methods. The smartphone scanning application was found to perform best overall when considering image quality and ease of use in the field. The automatic time-lapse camera was useful for measuring RER over several months without any human intervention. CONCLUSION: Our results show that inexpensive scanning and automated methods provide correct measurements of root elongation and length (but not diameter when using the time-lapse camera). These methods are capable of detecting fine roots to a diameter of 0.1 mm and can therefore be selected by the user depending on the data required.

Root biomass, turnover and net primary productivity of a coffee agroforestry system in Costa Rica: effects of soil depth, shade trees, distance to row and coffee age.

Background and Aims In Costa Rica, coffee (Coffea arabica) plants are often grown in agroforests. However, it is not known if shade-inducing trees reduce coffee plant biomass through root competition, and hence alter overall net primary productivity (NPP). We estimated biomass and NPP at the stand level, taking into account deep roots and the position of plants with regard to trees. Methods Stem growth and root biomass, turnover and decomposition were measured in mixed coffee/tree (Erythrina poeppigiana) plantations. Growth ring width and number at the stem base were estimated along with stem basal area on a range of plant sizes. Root biomass and fine root density were measured in trenches to a depth of 4 m. To take into account the below-ground heterogeneity of the agroforestry system, fine root turnover was measured by sequential soil coring (to a depth of 30 cm) over 1 year and at different locations (in full sun or under trees and in rows/inter-rows). Allometric relationships were used to calculate NPP of perennial components, which was then scaled up to the stand level. Key Results Annual ring width at the stem base increased up to 2·5 mm yr-1 with plant age (over a 44-year period). Nearly all (92 %) coffee root biomass was located in the top 1·5 m, and only 8 % from 1·5 m to a depth of 4 m. Perennial woody root biomass was 16 t ha-1 and NPP of perennial roots was 1·3 t ha-1 yr-1. Fine root biomass (0-30 cm) was two-fold higher in the row compared with between rows. Fine root biomass was 2·29 t ha-1 (12 % of total root biomass) and NPP of fine roots was 2·96 t ha-1 yr-1 (69 % of total root NPP). Fine root turnover was 1·3 yr-1 and lifespan was 0·8 years. Conclusions Coffee root systems comprised 49 % of the total plant biomass; such a high ratio is possibly a consequence of shoot pruning. There was no significant effect of trees on coffee fine root biomass, suggesting that coffee root systems are very competitive in the topsoil.

Root structure-function relationships in 74 species: evidence of a root economics spectrum related to carbon economy.

Although fine roots are important components of the global carbon cycle, there is limited understanding of root structure-function relationships among species. We determined whether root respiration rate and decomposability, two key processes driving carbon cycling but always studied separately, varied with root morphological and chemical traits, in a coordinated way that would demonstrate the existence of a root economics spectrum (RES). Twelve traits were measured on fine roots (diameter ≤ 2 mm) of 74 species (31 graminoids and 43 herbaceous and dwarf shrub eudicots) collected in three biomes. The findings of this study support the existence of a RES representing an axis of trait variation in which root respiration was positively correlated to nitrogen concentration and specific root length and negatively correlated to the root dry matter content, lignin : nitrogen ratio and the remaining mass after decomposition. This pattern of traits was highly consistent within graminoids but less consistent within eudicots, as a result of an uncoupling between decomposability and morphology, and of heterogeneity of individual roots of eudicots within the fine-root pool. The positive relationship found between root respiration and decomposability is essential for a better understanding of vegetation-soil feedbacks and for improving terrestrial biosphere models predicting the consequences of plant community changes for carbon cycling.

Seasonal Patterns of Fine Root Production and Turnover in a Mature Rubber Tree (Hevea brasiliensis Müll. Arg.) Stand- Differentiation with Soil Depth and Implications for Soil Carbon Stocks.

Fine root dynamics is a main driver of soil carbon stocks, particularly in tropical forests, yet major uncertainties still surround estimates of fine root production and turnover. This lack of knowledge is largely due to the fact that studying root dynamics in situ, particularly deep in the soil, remains highly challenging. We explored the interactions between fine root dynamics, soil depth, and rainfall in mature rubber trees (Hevea brasiliensis Müll. Arg.) exposed to sub-optimal edaphic and climatic conditions. A root observation access well was installed in northern Thailand to monitor root dynamics along a 4.5 m deep soil profile. Image-based measurements of root elongation and lifespan of individual roots were carried out at monthly intervals over 3 years. Soil depth was found to have a significant effect on root turnover. Surprisingly, root turnover increased with soil depth and root half-life was 16, 6-8, and only 4 months at 0.5, 1.0, 2.5, and 3.0 m deep, respectively (with the exception of roots at 4.5 m which had a half-life similar to that found between depths of 1.0 and 2.5 m). Within the first two meters of the soil profile, the highest rates of root emergence occurred about 3 months after the onset of the rainy season, while deeper in the soil, root emergence was not linked to the rainfall pattern. Root emergence was limited during leaf flushing (between March and May), particularly within the first two meters of the profile. Between soil depths of 0.5 and 2.0 m, root mortality appeared independent of variations in root emergence, but below 2.0 m, peaks in root emergence and death were synchronized. Shallow parts of the root system were more responsive to rainfall than their deeper counterparts. Increased root emergence in deep soil toward the onset of the dry season could correspond to a drought acclimation mechanism, with the relative importance of deep water capture increasing once rainfall ceased. The considerable soil depth regularly explored by fine roots, even though significantly less than in surface layers in terms of root length density and biomass, will impact strongly the evaluation of soil carbon stocks.

Modelling and predicting the spatial distribution of tree root density in heterogeneous forest ecosystems.

BACKGROUND AND AIMS: In mountain ecosystems, predicting root density in three dimensions (3-D) is highly challenging due to the spatial heterogeneity of forest communities. This study presents a simple and semi-mechanistic model, named ChaMRoots, that predicts root interception density (RID, number of roots m(-2)). ChaMRoots hypothesizes that RID at a given point is affected by the presence of roots from surrounding trees forming a polygon shape. METHODS: The model comprises three sub-models for predicting: (1) the spatial heterogeneity - RID of the finest roots in the top soil layer as a function of tree basal area at breast height, and the distance between the tree and a given point; (2) the diameter spectrum - the distribution of RID as a function of root diameter up to 50 mm thick; and (3) the vertical profile - the distribution of RID as a function of soil depth. The RID data used for fitting in the model were measured in two uneven-aged mountain forest ecosystems in the French Alps. These sites differ in tree density and species composition. KEY RESULTS: In general, the validation of each sub-model indicated that all sub-models of ChaMRoots had good fits. The model achieved a highly satisfactory compromise between the number of aerial input parameters and the fit to the observed data. CONCLUSIONS: The semi-mechanistic ChaMRoots model focuses on the spatial distribution of root density at the tree cluster scale, in contrast to the majority of published root models, which function at the level of the individual. Based on easy-to-measure characteristics, simple forest inventory protocols and three sub-models, it achieves a good compromise between the complexity of the case study area and that of the global model structure. ChaMRoots can be easily coupled with spatially explicit individual-based forest dynamics models and thus provides a highly transferable approach for modelling 3-D root spatial distribution in complex forest ecosystems.

A framework for identifying plant species to be used as 'ecological engineers' for fixing soil on unstable slopes.

Major reforestation programs have been initiated on hillsides prone to erosion and landslides in China, but no framework exists to guide managers in the choice of plant species. We developed such a framework based on the suitability of given plant traits for fixing soil on steep slopes in western Yunnan, China. We examined the utility of 55 native and exotic species with regard to the services they provided. We then chose nine species differing in life form. Plant root system architecture, root mechanical and physiological traits were then measured at two adjacent field sites. One site was highly unstable, with severe soil slippage and erosion. The second site had been replanted 8 years previously and appeared to be physically stable. How root traits differed between sites, season, depth in soil and distance from the plant stem were determined. Root system morphology was analysed by considering architectural traits (root angle, depth, diameter and volume) both up- and downslope. Significant differences between all factors were found, depending on species. We estimated the most useful architectural and mechanical traits for physically fixing soil in place. We then combined these results with those concerning root physiological traits, which were used as a proxy for root metabolic activity. Scores were assigned to each species based on traits. No one species possessed a suite of highly desirable traits, therefore mixtures of species should be used on vulnerable slopes. We also propose a conceptual model describing how to position plants on an unstable site, based on root system traits.

Deep phenotyping of coarse root architecture in R. pseudoacacia reveals that tree root system plasticity is confined within its architectural model.

This study aims at assessing the influence of slope angle and multi-directional flexing and their interaction on the root architecture of Robinia pseudoacacia seedlings, with a particular focus on architectural model and trait plasticity. 36 trees were grown from seed in containers inclined at 0° (control) or 45° (slope) in a glasshouse. The shoots of half the plants were gently flexed for 5 minutes a day. After 6 months, root systems were excavated and digitized in 3D, and biomass measured. Over 100 root architectural traits were determined. Both slope and flexing increased significantly plant size. Non-flexed trees on 45° slopes developed shallow roots which were largely aligned perpendicular to the slope. Compared to the controls, flexed trees on 0° slopes possessed a shorter and thicker taproot held in place by regularly distributed long and thin lateral roots. Flexed trees on the 45° slope also developed a thick vertically aligned taproot, with more volume allocated to upslope surface lateral roots, due to the greater soil volume uphill. We show that there is an inherent root system architectural model, but that a certain number of traits are highly plastic. This plasticity will permit root architectural design to be modified depending on external mechanical signals perceived by young trees.

Linking carbon supply to root cell-wall chemistry and mechanics at high altitudes in Abies georgei.

BACKGROUND AND AIMS: The mobile carbon supply to different compartments of a tree is affected by climate, but its impact on cell-wall chemistry and mechanics remains unknown. To understand better the variability in root growth and biomechanics in mountain forests subjected to substrate mass movement, we investigated root chemical and mechanical properties of mature Abies georgei var. smithii (Smith fir) growing at different elevations on the Tibet-Qinghai Plateau. METHODS: Thin and fine roots (0·1-4·0 mm in diameter) were sampled at three different elevations (3480, 3900 and 4330 m, the last corresponding to the treeline). Tensile resistance of roots of different diameter classes was measured along with holocellulose and non-structural carbon (NSC) content. KEY RESULTS: The mean force necessary to break roots in tension decreased significantly with increasing altitude and was attributed to a decrease in holocellulose content. Holocellulose was significantly lower in roots at the treeline (29·5 ± 1·3 %) compared with those at 3480 m (39·1 ± 1·0 %). Roots also differed significantly in NSC, with 35·6 ± 4·1 mg g(-1) dry mass of mean total soluble sugars in roots at 3480 m and 18·8 ± 2·1 mg g(-1) dry mass in roots at the treeline. CONCLUSIONS: Root mechanical resistance, holocellulose and NSC content all decreased with increasing altitude. Holocellulose is made up principally of cellulose, the biosynthesis of which depends largely on NSC supply. Plants synthesize cellulose when conditions are optimal and NSC is not limiting. Thus, cellulose synthesis in the thin and fine roots measured in our study is probably not a priority in mature trees growing at very high altitudes, where climatic factors will be limiting for growth. Root NSC stocks at the treeline may be depleted through over-demand for carbon supply due to increased fine root production or winter root growth.

(Not) Keeping the stem straight: a proteomic analysis of maritime pine seedlings undergoing phototropism and gravitropism.

BACKGROUND: Plants are subjected to continuous stimuli from the environment and have evolved an ability to respond through various growth and development processes. Phototropism and gravitropism responses enable the plant to reorient with regard to light and gravity. RESULTS: We quantified the speed of maritime pine seedlings to reorient with regard to light and gravity over 22 days. Seedlings were inclined at 15, 30 and 45 degrees with vertical plants as controls. A lateral light source illuminated the plants and stem movement over time was recorded. Depending on the initial angle of stem lean, the apical response to the lateral light source differed. In control and 15° inclined plants, the apex turned directly towards the light source after only 2 h. In plants inclined at 30° and 45°, the apex first reoriented in the vertical plane after 2 h, then turned towards the light source after 24 h. Two-dimensional gel electrophoresis coupled with mass spectrometry was then used to describe the molecular response of stem bending involved in photo- and gravi-tropism after 22 hr and 8 days of treatment. A total of 486 spots were quantitatively analyzed using image analysis software. Significant changes were determined in the protein accumulation of 68 protein spots. Early response gravitropic associated proteins were identified, which are known to function in energy related and primary metabolism. A group of thirty eight proteins were found to be involved in primary metabolism and energy related metabolic pathways. Degradation of Rubisco was implicated in some protein shifts. CONCLUSIONS: Our study demonstrates a rapid gravitropic response in apices of maritime pine seedlings inclined >30°. Little or no response was observed at the stem bases of the same plants. The primary gravitropic response is concomitant with a modification of the proteome, consisting of an over accumulation of energy and metabolism associated proteins, which may allow the stem to reorient rapidly after bending.

Plant growth modelling and applications: the increasing importance of plant architecture in growth models.

BACKGROUND: Modelling plant growth allows us to test hypotheses and carry out virtual experiments concerning plant growth processes that could otherwise take years in field conditions. The visualization of growth simulations allows us to see directly and vividly the outcome of a given model and provides us with an instructive tool useful for agronomists and foresters, as well as for teaching. Functional-structural (FS) plant growth models are nowadays particularly important for integrating biological processes with environmental conditions in 3-D virtual plants, and provide the basis for more advanced research in plant sciences. SCOPE: In this viewpoint paper, we ask the following questions. Are we modelling the correct processes that drive plant growth, and is growth driven mostly by sink or source activity? In current models, is the importance of soil resources (nutrients, water, temperature and their interaction with meristematic activity) considered adequately? Do classic models account for architectural adjustment as well as integrating the fundamental principles of development? Whilst answering these questions with the available data in the literature, we put forward the opinion that plant architecture and sink activity must be pushed to the centre of plant growth models. In natural conditions, sinks will more often drive growth than source activity, because sink activity is often controlled by finite soil resources or developmental constraints. PMA06: This viewpoint paper also serves as an introduction to this Special Issue devoted to plant growth modelling, which includes new research covering areas stretching from cell growth to biomechanics. All papers were presented at the Second International Symposium on Plant Growth Modeling, Simulation, Visualization and Applications (PMA06), held in Beijing, China, from 13-17 November, 2006. Although a large number of papers are devoted to FS models of agricultural and forest crop species, physiological and genetic processes have recently been included and point the way to a new direction in plant modelling research.

Understanding the impact of root morphology on overturning mechanisms: a modelling approach.

BACKGROUND AND AIMS: The Finite Element Method (FEM) has been used in recent years to simulate overturning processes in trees. This study aimed at using FEM to determine the role of individual roots in tree anchorage with regard to different rooting patterns, and to estimate stress distribution in the soil and roots during overturning. METHODS: The FEM was used to carry out 2-D simulations of tree uprooting in saturated soft clay and loamy sand-like soil. The anchorage model consisted of a root system embedded in a soil block. Two root patterns were used and individual roots removed to determine their contribution to anchorage. KEY RESULTS: In clay-like soil the size of the root-soil plate formed during overturning was defined by the longest roots. Consequently, all other roots localized within this plate had no influence on anchorage strength. In sand-like soil, removing individual root elements altered anchorage resistance. This result was due to a modification of the shape and size of the root-soil plate, as well as the location of the rotation axis. The tap root and deeper roots had more influence on overturning resistance in sand-like soil compared with clay-like soil. Mechanical stresses were higher in the most superficial roots and also in leeward roots in sand-like soil. The relative difference in stresses between the upper and lower sides of lateral roots was sensitive to root insertion angle. Assuming that root eccentricity is a response to mechanical stresses, these results explain why eccentricity differs depending on root architecture. CONCLUSIONS: A simple 2-D Finite Element model was developed to better understand the mechanisms involved during tree overturning. It has been shown how root system morphology and soil mechanical properties can modify the shape of the root plate slip surface as well as the position of the rotation axis, which are major components of tree anchorage.

Using three-dimensional plant root architecture in models of shallow-slope stability.

BACKGROUND: The contribution of vegetation to shallow-slope stability is of major importance in landslide-prone regions. However, existing slope stability models use only limited plant root architectural parameters. This study aims to provide a chain of tools useful for determining the contribution of tree roots to soil reinforcement. METHODS: Three-dimensional digitizing in situ was used to obtain accurate root system architecture data for mature Quercus alba in two forest stands. These data were used as input to tools developed, which analyse the spatial position of roots, topology and geometry. The contribution of roots to soil reinforcement was determined by calculating additional soil cohesion using the limit equilibrium model, and the factor of safety (FOS) using an existing slope stability model, Slip4Ex. KEY RESULTS: Existing models may incorrectly estimate the additional soil cohesion provided by roots, as the spatial position of roots crossing the potential slip surface is usually not taken into account. However, most soil reinforcement by roots occurs close to the tree stem and is negligible at a distance >1.0 m from the tree, and therefore global values of FOS for a slope do not take into account local slippage along the slope. CONCLUSIONS: Within a forest stand on a landslide-prone slope, soil fixation by roots can be minimal between uniform rows of trees, leading to local soil slippage. Therefore, staggered rows of trees would improve overall slope stability, as trees would arrest the downward movement of soil. The chain of tools consisting of both software (free for non-commercial use) and functions available from the first author will enable a more accurate description and use of root architectural parameters in standard slope stability analyses.