Deconstructing the root system of grasses through an exploration of development, anatomy and function.

ABSTRACTThe rice root system consists of three types of roots; main root, L-type and S-type lateral root (LR). These component roots are morphologically and histologically different, which is termed as heterorhizy. Root system hydraulic architecture is related to the unique features of component roots. We hypothesized that each component root contributes in different degrees to water uptake of the whole root system. Rice varieties IRAT 109 and Taichung 65 were grown in pots filled with soil under continuous waterlogged (CWL) and drought (CD) conditions until two weeks after heading. Morphology and histological structures of roots, which may regulate radial water movement, were compared among the three component roots. Moreover, hydraulic conductivity (Lpr) of the root system, which represents the water uptake ability, were measured with a pressure chamber. Based on a model that Lpr of the whole root system is a product of Lpr of each of the component roots and their surface areas, we found that the differences in Lpr between the two varieties and the plants grown under different soil water conditions for any of the component roots did not support the corresponding differences in the measured Lpr of the whole root system. In contrast, a significant and positive correlation was found between Lpr of the whole root system and the percentage of surface area of S-type LR but not for the other component roots. These results indicate S-type LR might have a higher contribution to Lpr of the whole root system than the other component roots.

Determining the variation in roots traits within a grass root system is important for understanding the role of fine roots in carbon and nutrient cycling in grassland ecosystems, where the majority of biomass and litter accumulation occur belowground. However, few studies have been conducted in this regard. In this study, the structural and chemical traits of shoot-derived and root-derived roots were examined in three perennial grasses—Cleistogenes squarrosa, Achnatherum sibiricum and Stipa grandis—aiming to explore structural differences, responses to nitrogen and water addition in different types of roots and their correlations with aboveground plant nitrogen. Our results showed significant differences between these two root types, with root-derived roots having higher N concentration, tissue density, and specific root length, but lower C: N and diameter than shoot-derived roots. Trait relationships between root N concentration and tissue density for the two root types differed from that reported among species. These traits in different types of roots were insensitive to resource addition. Furthermore, N concentration in shoot-derived roots was more strongly linked to aboveground plant N concentration than root-derived roots. The results of this study demonstrate structural differences within the root system that may reflect functional heterogeneity in grass roots.

The nodal root system is critical for the development of the mature root system in maize (Zea mays L.) and other grasses. Under drought conditions, nodal root axes may need to grow through surface soil that is dry, hard, and hot. These roots are known to have a superior ability to continue elongation at low water potentials relative to other organs of the plant, but the physiology of this response has been little studied. The objective of this study was to develop an experimental system that models the field situation in which upper soil layers dry, to enable studies of nodal root growth regulation under water deficit conditions. A divided-chamber experimental system was developed to allow the growth of maize primary and seminal root systems in well-watered conditions while the nodal root system is exposed to precise conditions of low soil water potential. The divided-chamber system was used to characterize nodal root growth responses to a range of soil water potentials under steady-state and reproducible conditions. Two contrasting genotypes, selected for differences in root growth response to water stress based on a previous study of the primary root, displayed similarly sensitive growth responses to -0.3 MPa soil, but different capacities to maintain high root tip water potential corresponding with different growth responses at lower soil water potentials. Both genotypes maintained relatively high nodal root tip water potentials in -2.0 MPa soil, despite the decreased soil water potential, suggesting a stress-induced response that enhances water transport to the root tip. The difference in high tissue water potential maintenance was seen not only between the contrasting genotypes but also between the first two developmental nodes of roots. The divided-chamber system provides a powerful experimental approach to investigate the physiological mechanisms regulating nodal root growth responses to adverse soil conditions. Future studies may include measurements of hydraulic conductivity, anatomical characterization of vascular elements near the growth zone, aquaporin content and activity, and suberin deposition in response to low soil water potentials.

Wheat was grown in a slightly acid grey sand at six levels of phosphorus and four levels of nitrogen, in the presence and absence of inoculum of Gaeurnannomyces graminis tritici (Ggt) in a glasshouse experiment. Adequate nutrition of N (400 mg/pot) and P (100 mg/pot) for wheat plants was required to overcome take-all of wheat grown for 46 days. Severely N and P deficient plants had 60% of their seminal and nodal roots infected by take-all. As the plants responded to increasing levels of N and P, the percentage of infected nodal and seminal roots steadily declined to 0% and 10% for each root system respectively at luxury levels of P and adequate N. In both the seminal and nodal root system, increasing the N and P supply decreased the length of proximal lesions (closest to seed) and increased the length of the root between the crown and the proximal lesion. The seminal root system was more severely infected with take-all than the nodal root system.

Root systems of woody plants generally display a strong relationship between the cross-sectional area or cross-sectional diameter (CSD) of a root and the dry weight of biomass (DWd) or root volume (Vd) that has grown (i.e., is descendent) from a point. Specification of this relationship allows one to quantify root architectural patterns and estimate the amount of material lost when root systems are extracted from the soil. However, specifications of this relationship generally do not account for the fact that root systems are comprised of multiple types of roots. We assessed whether the relationship between CSD and Vd varies as a function of root type. Additionally, we sought to identify a more accurate and time-efficient method for estimating missing root volume than is currently available. We used a database that described the 3D root architecture of Pinus pinaster root systems (5, 12, or 19 years) from a stand in southwest France. We determined the relationship between CSD and Vd for 10,000 root segments from intact root branches. Models were specified that did and did not account for root type. The relationships were then applied to the diameters of 11,000 broken root ends to estimate the volume of missing roots. CSD was nearly linearly related to the square root of Vd, but the slope of the curve varied greatly as a function of root type. Sinkers and deep roots tapered rapidly, as they were limited by available soil depth. Distal shallow roots tapered gradually, as they were less limited spatially. We estimated that younger trees lost an average of 17% of root volume when excavated, while older trees lost 4%. Missing volumes were smallest in the central parts of root systems and largest in distal shallow roots. The slopes of the curves for each root type are synthetic parameters that account for differentiation due to genetics, soil properties, or mechanical stimuli. Accounting for this differentiation is critical to estimating root loss accurately.

The patterns of ring width throughout the root systems and stems of several 32- to 36-year-old, plantation-grown red pine were analyzed. All root systems showed a basic similarity in their radial sequences of ring width. Within a root system, individual sequences ranged from those similar to the radial sequence at the stem base to ones apparently dissimilar. At the stem–root base, the increasing distance to the tree crown with time and changes in mechanical stresses with increasing tree size appeared to play major roles in the development of the basic pattern. The position and type of root, the environment of its apices, and changes in the transportation pathways in the xylem and phloem at branching points may all have influenced the pattern in individual roots. The pattern of deposition of xylem at branching points was visualized as an interaction between the stream of growth regulators and assimilates towards the root tips and the stream of xylem water (and (or) dissolved substances) towards the root base. Discontinuities in the growth layers did not occur in the stem but increased with time in the roots, particularly in the surface horizontal roots. Moisture availability modified the radial sequence patterns and probably influenced the shifts in longitudinal distribution of radial increment from year to year within and between the stem and the root system. The maximum current annual increment in root wood volume occurred before that in the stem.

Root architecture can be very important in plant productivity. The importance of studies on root morphology and development is discussed to improve seedling growth. Root systems of dicotyledonous species are reviewed, with emphasis on differences between growth of basal and lateral roots. The presence of different types of roots in plant species suggests possible differences in function as well. The architecture of a root system related to its functions is considered. Classical methods for studying root systems comprise excavation of root system, direct observation, and indirect analyses. While the first method is destructive and the third is effective in understanding root architecture only on a relatively gross scale, observation methods allow the scientist a complete a nondestructive architectural study of a root system. The three groups are reviewed related to their potential to give valuable information related to the root architecture and development of the seedling, with emphasis on the availability of a medium-transparent plant-growing system, enabling nondestructive daily observations and plant measurements under controlled environmental conditions. Effects of CO 2 enrichment on seedling growth is reviewed, emphasizing the effects of CO 2 on root growth.

Large river floodplains are convenient model systems to test for variation in animal and plant community structure, as they have a variety of habitats and substrates and are generally dynamic systems through the occurrence of flood pulses with varying intensity. South American floodplain systems furthermore have unique types of substrates, in the form of root systems of floating macrophytes. Here, we investigate the variation in ostracod (small, bivalved crustaceans) communities in relation to substrates and related environmental variables. Sampling was effected in 2004 in the alluvial valley of the upper Parana River, Brazil, in the wet and dry seasons. Five different substrates, including littoral sediment and four macrophyte species root and leaf systems, in four hydrological systems and a variety of habitat types, were sampled. Fifty-four species of Ostracoda were found. Variation partitioning analysis (RDA) showed that ostracod communities significantly differed between different substrates, mainly between the littoral and plants with small root systems (Eichhornia azurea) on the one hand, and plants with large and complex root systems on the other hand (Eichhornia crassipes and Pistia stratiotes). RDA analyses indicated that the pleuston (biotic communities associated with root systems of floating plants) of E. crassipes comprised more non-swimming species than the pleuston of the smaller roots of P. stratiotes, but species-level Kruskal–Wallis analyses could not detect significant differences between both macrophyte species. Also habitat type and hydrological systems contributed to variation amongst ostracod communities, but less so than the factor substrate. Abiotic factors also contributed to variation, but the ranges of all measured water chemistry variables were narrow. This uniformity in abiotic factors, which might be owing to the occurrence of large flooding events, unites all water bodies, even those that are generally separated.

In numerous studies dealing with roots of woody plants, a description of the root system architecture is needed. During the twentieth century, several manual measurement methods were used, depending on the objectives of study. Due to the difficulties in accessing the roots and the duration of measurements, the studies generally involved a low number of root systems, were often qualitative and focused only on one specific application. Quantitative methods in plant architecture were largely developed in the last 40 years for aerial architecture. However, root systems have particular features and often need specific procedures. Since the end of the 1990s, new devices and techniques have been available for coarse root architecture measurements including volume location techniques (non-invasive or destructive) and manual or semi-automatic 3D digitising. Full 3D root system architecture dynamics was also reconstructed from partial measurements using modelling procedures. On the one hand, non-invasive and automatic techniques need more development to obtain full 3D architecture, i.e. geometry and topology. On the other hand, both one inexpensive manual and one semi-automatic digitizing procedure are now available to measure precisely and rapidly the full 3D architecture of uprooted and excavated coarse root systems. Specific software and a large number of functions are also available for an in-depth analysis of root architecture and have already been used in a dozen of research papers including a fairly large sample of mature trees. A comprehensive analysis of root architecture can be achieved by classifying individual roots in several root types through architectural analysis. The objective of this paper is both to give a detailed overview of the state of the art techniques for 3D root system architecture measurement and analysis and to give examples of applications in this field. Practical details are also given so that this paper can be used as a sort of manual for people who want to improve their practice or to enter this quite new research field.

The perfect storm. That is the term used by Gerald C. Nelson to describe the triple challenge of increasing food production while adapting to climate change and reducing the environmental impact of agricultural systems. Nowadays, conventional farming systems are showing some limitations, such as low resources use efficiency and poor ecosystems services that appear to be associated to the loss of plant diversity and perenniality in crop rotations. In addition, water, the most important yield limiting factor worldwide, will increasingly restrict food production in the future due to rainfall shortage and increase in human consumption. In such context, perennial crops, with denser and deeper root system could use resources in deep soil layers that are logically inaccessible to crops with shallower root system. The goal of this thesis was therefore to investigate the root growth and water uptake capacity of intermediate wheatgrass (Kernza®) and alfalfa, two deep rooted perennial crops, under field conditions and at great soil depth (i.e. 1.0-2.5 m).
\n\tMaintaining hydraulic continuity along the soil-plant-atmosphere continuum is a prerequisite for deep water uptake. At the plant level, hydraulic conductivity depends on complex anatomical and physiological processes among which the root system constitutes the second largest resistance to water flow. Therefore, in depth characterisation of root and xylem anatomy was done to understand the hydraulic properties of the crop root systems, with a focus on their evolution with soil depth. Crops were grown in the field, rhizoboxes, mesocosms and solution culture to take into account the variability of root type and soil depth as well as growing environment. For both crops, axial hydraulic conductance decreased with soil depth and along individual root segment. Alfalfa roots had greater axial hydraulic conductance in comparison to intermediate wheatgrass roots, especially at depth. Root and xylem anatomy were highly variable across crop species, root types and growing environments. In parallel, a combination of imaging and sensor technology, stable isotope techniques and a modelling approach was used to study root growth and water uptake under field conditions during the 2018-2019 seasons. Both crops presented roots down to 2.0 m soil depth that were active in terms of water uptake. Alfalfa had greater root length at depth and absorbed twice as much water below 1 m soil depth, than intermediate wheatgrass. For both crops, model simulations predicted that water uptake in deep soil layers (i.e. 1.5 – 2.0 m) increase (i.e.>30%) under dry conditions.
\n\tThis thesis brings insights into the understudied field of root growth and water uptake at great soil depth. Particular efforts were put in understanding the environmental and agricultural contexts in which deep root growth, deep water uptake and the development of perennial cropping systems would be possible and favourable.

In water-limited ecosystems such as the shortgrass steppe of North America, the ability of plants to absorb sufficient water for growth and reproduction is dependent upon the characteristics of their root systems. Our objective was to evaluate the spatial dis- tributions of roots of three plant life forms: a grass (Bouteloua gracilis) (H.B.K.) Lag., a dwarf shrub (Gutierrezia sarothrae (Pursh.) Britt. & Rusby) and a shrub (Atriplex canescens (Pursh.) Nutt.). Our interests were in a qualitative evaluation of evidence that different life forms obtained water from different soil layers. Root systems were excavated and evaluated in terms of horizontal and vertical extension, and proportion of total root length in each 10-cm layer of the soil profile. Bouteloua gracilis had over 80% of its roots in the top 50 cm of the soil, whereas the depth at which Atriplex canescens had over 80% of its roots was 100 cm. The corresponding depth for Gutierrezia sarothrae was 80 cm. Both of the woody plants had deeper and more widely spreading root systems than the grass. The environment of the shortgrass steppe is characterized by low and variable water avail- ability. These conditions lead to a distribution of available water that is biased toward the upper soil layers. The greater concentration of roots of Bouteloua gracilis in the soil layers near the surface compared to the two woody species provides a potential explanation for its dominance in the shortgrass steppe.

SPACSYS: Integration of a 3D root architecture component to carbon, nitrogen and water cycling—Model description

OPTIMALIZATION OF OXYGEN LEVELS IN ROOT SYSTEMS AS EFFECTIVE CULTIVATION TOOL

A developmentally based categorization of branching in Trifolium repens L.: influence of nodal roots.

In this talk we introduce generalised Calogero–Moser models and demonstrate their integrability by constructing universal Lax pair operators. These include models based on non-crystallographic root systems, that is the root systems of the finite reflection groups, H 3, H 4, and the dihedral group I 2(m), besides the well-known ones based on crystallographic root systems, namely those associated with Lie algebras. Universal Lax pair operators for all of the generalised Calogero–Moser models and for any choices of the potentials are linear combinations of the reflection operators. The equivalence of the Lax pair with the equations of motion is proved by decomposing the root system into a sum of two-dimensional sub-root systems, A 2, B 2, G 2, and I 2(m). The root type and the minimal type Lax pairs, derived in our previous papers, are given as the simplest representations. The spectral parameter dependence plays an important role in the Lax pair operators, which bear a strong resemblance to the Dunkl operators.