Unravelling the response of poplar (Populus nigra) roots to mechanical stress imposed by bending

Abstract

Mechanical stress is a widespread environmental condition that can be caused by several factors (i.e. gravity, touch, wind, soil density, soil compaction and grazing, slope) and that can severely affect plant stability. In response to mechanical stress and to improve their anchorage, plants have developed complex mechanisms to detect mechanical perturbation and to induce a suite of modifications at anatomical, physiological, biochemical, biophysical and molecular level. Although it is well recognized that one of the primary functions of root systems is to anchor the plant to the soil, root response to mechanical stresses have been investigated mainly at morphological and biomechanical level, whereas investigations about the molecular mechanisms underlying these important alterations are still in an initial stage. We have used an experimental system in which the taproot poplar seedlings are bent to simulate mechanical perturbation to begin investigate the mechanisms involved in root response to mechanical stress. The results reported herein show that, in response to bending, the poplar root changes its morphology by emitting new lateral roots, and its biomechanical properties by increasing the root biomass and lignin synthesis. In addition, using a proteomic approach, we found that several proteins involved in the signal transduction pathway, detoxification and metabolism are up-regulated and/or down-regulated in the bent root. These results provide new insight into the obscure field of woody root response to mechanical stress, and can serve as a basis for future investigations aimed at unravelling the complex mechanism involved in the reaction of root biology to environmental stress.     

Improving vigour assessment of pine (Pinus nigra Arnold) seedlings before their use in reforestation

ABSTRACT

We investigated whether changes in the root system of pine seedlings induced by stress (lifting of bare-root seedlings from the nursery bed irrespective of dormancy; prolonged storage of bare-root seedlings in a cold room) could provide a measure of plant vigour. Physiological parameters, such as growth potential and root electrolyte leakage, and morphological parameters, such as root length and number of root tips, were calculated. Computerised image analysis was used to measure root growth, overall and based on root-diameter class (0–0.5 mm, 0.5–1.0 mm and 1.0–1.5 mm). The efficiency of vigour assessment was evaluated by correlating the data for each parameter with percentage seedling survival. Root growth potential was more efficient than root electrolyte leakage, but both parameters were affected by seedling age. Total root length was a more efficient indicator of plant vigour than root tip number, particularly when referred to roots of the same diameter class. A comparative analysis of physiological and morphological parameters referred to the root systems improves their relative effciency.     

Modification of root architecture in woody plants is possible for the presence of two different mechanisms of lateral root production: The effect of slope in Spartium junceum L. seedlings

Abstract

This paper investigates the modification of root architecture of Spartium junceum L. seedlings grown in slope condition. It is reported that 50% of the total number of lateral roots are concentrated in few centimetres of the taproot near the collar. The anatomical analysis of transverse sections along the taproot axis reveals that this taproot zone is characterised by two types of lateral roots: one with a trace extending to the centre of the vascular cylinder by following the path of a medullar ray; one with a trace which ends in the vascular cambium. The first type may be lateral roots originated from the taproot primary structure; the second type seems to be lateral roots developing later when a secondary structure has completely substituted the primary structure. The emission of this second type of lateral roots seems to be strongly controlled by environmental conditions with considerable consequences upon the overall root architecture. In the example reported in this paper, young plants growing under mechanical stress due to a slope develop asymmetric root architecture with lateral roots elongating in two prevalent directions: up-slope or down-slope. This asymmetric architecture is produced in the zone of the taproot where a secondary structure is present and represents the plant response to the need of increasing its anchorage strength.     

Root architecture and hydraulic conductance in nutrient deprived Pistacia lentiscus L. seedlings

Plants respond to low nutrient availability by modifying root morphology and root system topology. Root responses to nitrogen (N) and phosphorus (P) limitation may affect plant capacity to withstand water stress. But studies on the effect of nutrient availability on plant ability to uptake and transport water are scarce. In this study, we assess the effect of nitrogen and phosphorus limitation on root morphology and root system topology in Pistacia lentiscus L seedlings, a common Mediterranean shrub, and relate these changes to hydraulic conductivity of the whole root system. Nitrogen and phosphorus deprivation had no effect on root biomass, but root systems were more branched in nutrient limited seedlings. Total root length was higher in seedlings subjected to phosphorus deprivation. Root hydraulic conductance decreased in nutrient-deprived seedlings, and was related to the number of root junctions but not to other architectural traits. Our study shows that changes in nutrient availability affect seedling water use by modifying root architecture. Changes in nutrient availability should be taken into account when evaluating seedling response to drought.     

The influence of slope on <Emphasis Type="Italic">Spartium junceum</Emphasis> root system: morphological,anatomical and biomechanical adaptation

Root systems have a pivotal role in plant anchorage and their mechanical interactions with the soil may contribute to soil reinforcement and stabilization of slide-prone slopes. In order to understand the responses of root system to mechanical stress induced by slope, samples of Spartium junceum L., growing in slope and in plane natural conditions, were compared in their morphology, biomechanical properties and anatomical features. Soils sampled in slope and plane revealed similar characteristics, with the exception of organic matter content and penetrometer resistance, both higher in slope. Slope significantly influenced root morphology and in particular the distribution of lateral roots along the soil depth. Indeed, first-order lateral roots of plants growing on slope condition showed an asymmetric distribution between up- and down-slope. Contrarily, this asymmetric distribution was not observed in plants growing in plane. The tensile strength was higher in lateral roots growing up-slope and in plane conditions than in those growing down-slope. Anatomical investigations revealed that, while roots grown up-slope had higher area covered by xylem fibers, the ratio of xylem and phloem fibers to root diameter did not differ among the three conditions, as also, no differences were found for xylem fiber cell wall thickness. Roots growing up-slope were the main contributors to anchorage properties, which included higher strength and higher number of fibers in the xylematic tissues. Results suggested that a combination of root-specific morphological, anatomical and biomechanical traits, determines anchorage functions in slope conditions.     

Arabidopsis Lateral Root Development 3 is essential for early phloem development and function, and hence for normal root system development

We have identified a gene, Lateral Root Development 3 (LRD3), that is important for maintaining a balance between primary and lateral root growth. The lrd3 mutant has decreased primary root growth and increased lateral root growth. We determined that the LRD3 gene encodes a LIM-domain protein of unknown function. LRD3 is expressed only in the phloem companion cells, which suggested a role in phloem function. Indeed, while phloem loading and export from the shoot appear to be normal, delivery of phloem to the primary root tip is limited severely in young seedlings. Abnormalities in phloem morphology in these seedlings indicate that LRD3 is essential for correct early phloem development. There is a subsequent spontaneous recovery of normal phloem morphology, which is correlated tightly with increased phloem delivery and growth of the primary root. The LRD3 gene is one of very few genes described to affect phloem development, and the only one that is specific to early phloem development. Continuous growth on auxin also leads to recovery of phloem development and function in lrd3, which demonstrates that auxin plays a key role in early phloem development. The root system architecture and the pattern of phloem allocation in the lrd3 root system suggested that there may be regulated mechanisms for selectively supporting certain lateral roots when the primary root is compromised. Therefore, this study provides new insights into phloem-mediated resource allocation and its effects on plant root system architecture.     

樟树(Cinnamomum camphora)幼苗细根形态对氮磷添加和幼苗密度的响应

全球氮沉降对森林生态系统结构和功能的影响已成为现代生态学研究热点之一,我国华南地区氮沉降的增长引起了土壤酸化和磷限制加剧等一系列生态问题。密度制约着植物个体对环境资源的吸收利用,是自然界中十分重要的选择压力之一。因此研究樟树(Cinnamomum camphora)幼苗的细根形态对氮磷添加和密度的响应,有利于了解亚热带树木根系对氮沉降和磷添加与林分密度的响应过程和机制,并为全球变化背景下樟树林生态系统的管理提供依据。本研究以1年生樟树幼苗为试验材料,选择氯化铵(NH_4Cl)作为氮肥以模拟大气氮沉降,并且以二水合磷酸二氢钠(NaH_2PO_4·2H_2O)模拟磷添加,氮磷处理设置4个水平,即对照、施N、施P和施N+P;种植密度设置10、20、40和80株/m~2 4个水平。测定各处理樟树幼苗细根的根长、表面积、体积和根尖数,分析氮磷添加、密度和两者交互作用对樟树幼苗细根的影响。研究结果表明,与对照处理相比,N、P和N+P处理促进了幼苗细根长度、表面积、体积以及根尖数的增加。低密度条件下的N添加对幼苗根系形态的促进效果强于P添加。N+P处理对10、20、40株/m~2幼苗根系形态的促进效果最佳,而各处理对80株/m~2幼苗根系形态的促进效果均无显著性差异。随着种植密度的增大,幼苗细根长度、表面积、体积和根尖数均减少。樟树幼苗的细根长度、表面积、体积和根尖数在各密度间和不同氮磷添加处理间均有显著性差异,密度和氮磷处理间的交互作用对根系形态各指标均无显著影响。     

垂直方向磷素竞争对杉木根系生长及生物量分配的影响

针对自然环境中有效磷养分主要分布于土壤表层而容易导致植物根系激烈竞争的问题,选择同一杉木(Cunninghamia lanceolata)无性系幼苗为研究对象,采用水平方向空间狭小而垂直方向空间大的室内盆栽模拟装置,以单株种植为对照,构建双株种植的竞争处理,通过设置3个供磷水平:不供磷处理(0 mg/kg KH_2PO_4)、低磷处理(6 mg/kg KH_2PO_4)和正常供磷处理(12 mg/kg KH_2PO_4),采用破坏性试验方式收获,分别在试验的前期(50 d)、中期(100 d)和后期(150 d)测定不同处理条件下杉木幼苗根系生物量与根系形态的变化,研究邻株杉木根系在垂直方向上对有限磷素资源的竞争策略。结果表明:竞争处理和供磷水平对杉木幼苗根系长度、平均直径等形态指标的影响存在交互作用(P0.05),对杉木幼苗生物量分配、比根长等指标的影响均不存在明显的交互作用(P0.05)。竞争处理中杉木根系形态增量均明显高于非竞争处理的单株幼苗,且随着胁迫时间的增加,根系形态增量均呈现显著的上升趋势,其中在胁迫中期和后期的增量明显高于前期,且邻株竞争处理明显提高了杉木的比根长,提升了根系觅磷的能力;随着供磷水平的提高,根表面积和根体积增量大体上呈现先上升后下降的趋势。与非竞争处理相比,竞争条件下杉木地上部生物积累量差异不明显,而根系生物量、根冠比均低于非竞争处理的单株幼苗。     

Root Development and Nutrient Uptake

Root system formation proceeds in close coordination with shoot growth. Accordingly, root growth and its functions are regulated tightly by the shoot through materials cycling between roots and shoots. A plant root system consists of different kinds of roots that differ in morphology and functions. The spatial configuration and distribution of these roots determine root system architecture in the soil, which in turn primarily regulates the acquisition of soil resources like nutrients and water. Morphological and physiological properties of each root and the concomitant tissues further affect nutrient uptake and transport, while the root traits that are related to such acquisition also depend on the kinds of nutrients and their mobility in the soil. In addition, mechanisms involved in the uptake and transport of mineral nutrients recently have been elucidated at the molecular level. A number of genes for acquisition and transport of various mineral nutrients have been identified in model plant systems such as Arabidopsis thaliana, and rice, and in other plant species. An integration of studies on nutrient behavior in soils and the morphological and physiological functions of root systems will further elucidate the mechanism of plant nutrient uptake and transport by roots, and offer a real possibility of genetically improving crop productivity in problem soils.

     

模拟增雨对荒漠植物幼苗生长和根系形态的影响

降雨是荒漠生态系统过程和功能的最重要限制因子,荒漠植物幼苗对生长季降雨的变化极端敏感。为探讨荒漠植物对未来降雨格局变化的响应,选取乌兰布和沙漠两种典型荒漠植物幼苗(白刺和油蒿)为研究对象,根据生长季内(6—9月)每次降雨量,进行不同梯度的人工模拟增雨试验(CK:自然降雨、A:增雨25%、B:增雨50%、C:增雨75%、D:增雨100%),研究两种植物幼苗生长和根系形态特征对降雨量变化的响应。结果表明:(1)不同增雨处理对白刺和油蒿幼苗的地上部生长有显著影响(P0.05),增雨处理的白刺和油蒿幼苗的株高、平均冠幅和基径显著高于CK,并随着增雨量的增大而增大(白刺基径除外);(2)增雨处理之间、白刺和油蒿之间在总根长、总表面积、平均直径、总体积、根尖数和分叉数均有显著差异(P0.05)。对白刺幼苗而言,B处理和C处理的根系参数均显著大于CK、A和D处理,且B和C处理之间没有显著差异(平均直径除外);对油蒿幼苗而言,随着增雨量的增加,油蒿总根长、总表面积、总体积、根尖数和分叉数呈现逐渐增加的趋势,而平均直径呈现先增加后降低的趋势,且在B处理下达到最大值。(3)增雨处理显著降低了白刺幼苗的根冠比,而对油蒿幼苗的根冠比没有显著影响,并且白刺幼苗根冠比显著大于油蒿幼苗。(4)白刺和油蒿幼苗的根系主要分布在0—0.5 mm和0.5—1 mm内,2 mm的细根比例分别在B和D处理下达到最大值。这表明白刺和油蒿幼苗能够通过调整地上部生长和根系形态来适应降雨的变化。     

植物根系耐盐机制的研究进展

植物根系能够摄取土壤环境中的养分与水分,在植物的生长发育中起重要的作用。植物根系由于直接与土壤环境相接触会受到非生物胁迫较大的影响。盐胁迫是主要的非生物胁迫之一,对植物根系会产生较大的伤害。综述根系在组织形态和细胞水平上对盐胁迫的应答,以及根系响应盐胁迫的信号传导途径、转录因子与基因,对植物根部耐盐机制的解析和植物耐盐基因工程工具基因的挖掘具有重要意义。     

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.     

隔沟交替灌溉条件下玉米根系形态性状及结构分布

为揭示根系对土壤环境的适应机制,研究了隔沟交替灌溉条件下玉米根系形态性状及结构分布。以垄位和坡位的玉米根系为研究对象,利用Minirhizotrons法研究了根系（活/死根）的长度、直径、体积、表面积、根尖数和径级变化及其与土壤水分、土温和水分利用效率（WUE）的相关关系。结果表明,对于活根,在坡位非灌水区域复水后根系平均直径减小,而根系日均生长速率、单位面积土壤根系体积密度、根尖数和表面积均增大,并随灌水区域土壤水分的消退逐渐减小;对于死根,在坡位非灌水区域复水后根系日均死亡速率、根系体积密度、根尖数和表面积变化均减小,其中根系死亡速率和死根直径随土壤水分的消退逐渐降低,而死根体积密度、根尖数和表面积分布随土壤水分降低呈增大趋势;在垄位,根系形态分布趋势与坡位一致,除根系直径与与坡位比较接近外,其他根系形态值均小于坡位。将根系分成4个径级区间分析根系的形态特征,结果表明在根系长度和体积密度分布中以2.5－4.5 mm径级的根系所占比例最大,在根尖数和根系表面积分布中以0.0－2.5 mm径级的根系为主。通过显著性相关分析,死根直径、体积密度、活根表面积等根系形态与土壤含水率、土壤温度和WUE间均存在显著或极显著的正相关关系,部分根系形态指标（如根系的生长速率、活根体积密度）只与坡位土壤含水量、土壤温度具有明显的相关性,表明隔沟交替灌溉对坡位根系形态的调控作用比垄位显著。     

A density-based approach for the modelling of root architecture: application to Maritime pine (Pinus pinaster Ait.) root systems

Root morphology influences strongly plant/soil interactions. However, the complexity of root architecture is a major barrier when analysing many phenomena, e.g. anchorage, water or nutrient uptake. Therefore, we have developed a new approach for the representation and modelling of root architecture based on branching density. A general root branching density in a space of finite dimension was used and enabled us to consider various morphological properties. A root system model was then constructed which minimizes the difference between measured and simulated root systems, expressed with functions which map root density in the soil. The model was tested in 2D using data from Maritime pine Pinus pinaster Ait. structural roots as input. We showed that simulated and real root systems had similar root distributions in terms of radial distance, depth, branching angle and branching order. These results indicate that general density functions are not only a powerful basis for constructing models of architecture, but can also be used to represent such structures when considering root/soil interaction. These models are particularly useful in that they provide a local morphological characterization which is aggregated in a given unit of soil volume.     

降雨格局变化对白刺幼苗根系形态特征的影响

为探讨荒漠植物白刺幼苗根系形态对降雨格局变化的响应特征,设置3个降雨量梯度(W-、W、W+)和2个降雨间隔时间梯度(T、T+)进行人工模拟试验,结果表明,1)降雨量和降雨间隔时间对白刺幼苗根系形态有不同程度的影响,且降雨量的作用效应更大。2)降雨量相同时,延长降雨间隔时间均使白刺幼苗主根长、根系平均直径、根体积和根表面积减小,但总根长和根系生物量和总生物量却增加,在高降雨量条件下(W+)延长降雨间隔时间白刺幼苗比根长和比表面积分别增加了45.09%和20.20%,但差异均不显著。3)降雨间隔时间相同时,降雨量减少30%仅使主根长平均增加12.06%,总根长、根平均直径、根体积和根表面积等根系形态指标均显著减少,比根长和比表面积变化不大;降雨量增加30%仅使比表面积显著增加,其余各形态指标差异均不显著,低降雨量条件下(W-)主根长与根冠比达到最大,其他指标均在高降雨量条件下(W+)达到最大。4)对8个根系形态参数进行主成分分析,根系生物量、总根长、总根表面积、比根长、比表面积和根体积6个根系生态参数受降雨格局影响显著。     

Snowpack,fire, and forest disturbance: interactions affect montane invasions by non‐native shrubs

Montane regions worldwide have experienced relatively low plant invasion rates, a trend attributed to increased climatic severity, low rates of disturbance, and reduced propagule pressure relative to lowlands. Manipulative experiments at elevations above the invasive range of non‐native species can clarify the relative contributions of these mechanisms to montane invasion resistance, yet such experiments are rare. Furthermore, global climate change and land use changes are expected to cause decreases in snowpack and increases in disturbance by fire and forest thinning in montane forests. We examined the importance of these factors in limiting montane invasions using a field transplant experiment above the invasive range of two non‐native lowland shrubs, Scotch broom (Cytisus scoparius) and Spanish broom (Spartium junceum), in the rain–snow transition zone of the Sierra Nevada of California. We tested the effects of canopy closure, prescribed fire, and winter snow depth on demographic transitions of each species. Establishment of both species was most likely at intermediate levels of canopy disturbance, but at this intermediate canopy level, snow depth had negative effects on winter survival of seedlings. We used matrix population models to show that an 86% reduction in winter snowfall would cause a 2.8‐fold increase in population growth rates in Scotch broom and a 3.5‐fold increase in Spanish broom. Fall prescribed fire increased germination rates, but decreased overall population growth rates by reducing plant survival. However, at longer fire return intervals, population recovery between fires is likely to keep growth rates high, especially under low snowpack conditions. Many treatment combinations had positive growth rates despite being above the current invasive range, indicating that propagule pressure, disturbance, and climate can all strongly affect plant invasions in montane regions. We conclude that projected reductions in winter snowpack and increases in forest disturbance are likely to increase the risk of invasion from lower elevations.     

Form matters: morphological aspects of lateral root development

Background

The crucial role of roots in plant nutrition, and consequently in plant productivity, is a strong motivation to study the growth and functioning of various aspects of the root system. Numerous studies on lateral roots, as a major determinant of the root system architecture, mostly focus on the physiological and molecular bases of developmental processes. Unfortunately, little attention is paid either to the morphological changes accompanying the formation of a lateral root or to morphological defects occurring in lateral root primordia. The latter are observed in some mutants and occasionally in wild-type plants, but may also result from application of external factors.

Scope and Conclusions

In this review various morphological aspects of lateral branching in roots are analysed. Morphological events occurring during the formation of a typical lateral root are described. This process involves dramatic changes in the geometry of the developing organ that at early stages are associated with oblique cell divisions, leading to breaking of the symmetry of the cell pattern. Several types of defects in the morphology of primordia are indicated and described. Computer simulations show that some of these defects may result from an unstable field of growth rates. Significant changes in both primary and lateral root morphology may also be a consequence of various mutations, some of which are auxin-related. Examples reported in the literature are considered. Finally, lateral root formation is discussed in terms of mechanics. In this approach the primordium is considered as a physical object undergoing deformation and is characterized by specific mechanical properties.     

Root/Shoot Allocation and Root Architecture in Seedlings: Variation among Forest Sites, Microhabitats, and Ecological Groups1

I analyzed patterns of variation in root mass allocation and root morphology among seedlings of woody species in relation to environmental factors in four Neotropical forests. Among forests, I explored the response of root traits to sites varying in water or nutrient availability. Within each forest, I explored the plastic response of species to different microhabitats: gaps and understory. Additionally, I explored evidence for life history correlation of root and shoot traits by comparing species differing in their successional group (light‐demanding [22 spp.] or shade tolerant [27 spp.]) and germination type (species with photosynthetic cotyledons or species with reserve cotyledons). At each forest site, young seedlings from 10 to 20 species were excavated. A total of 55 species was collected in understory conditions and 31 of them were also collected in gaps. From each seedling, six morphological ratios were determined. Allocation to roots was higher in forest sites with the lowest soil resources. Roots were finer and longer in the most infertile site, while roots were deeper in the site with the longest dry season. Seedling traits did not differ between germination types. Shade tolerant species allocated more to roots and developed thicker roots than light‐demanding species. Light‐demanding species showed stronger plastic responses to habitat than shade tolerant species, and species with photo‐synthetic cotyledons showed lower plasticity than species with reserve cotyledons. Overall, these results suggest that among Neotropical species, root allocation and root morphology of seedlings reflect plant adjustments to water or nutrient availability at geographic and microhabitat scales. In addition, life history specialization to light environments is suggested by differences among groups of species in their allocation to roots and in their root morphology.     

Differential expression profiles of growth-related genes in the elongation zone of maize primary roots

Growth in the apical elongation zone of plant roots is central to the development of functional root systems. Rates of root segmental elongation change from accelerating to decelerating as cell development proceeds from newly formed to fully elongated status. One of the primary variables regulating these changes in elongation rates is the extensibility of the elongating cell walls. To help decipher the complex molecular mechanisms involved in spatially variable root growth, we performed a gene identification study along primary root tips of maize (Zea mays) seedlings using suppression subtractive hybridization (SSH) and candidate gene approaches. Using SSH we isolated 150 non-redundant cDNA clones representing root growth-related genes (RGGs) that were preferentially expressed in the elongation zone. Differential expression patterns were revealed by Northern blot analysis for 41 of the identified genes and several candidate genes. Many of the genes have not been previously reported to be involved in root growth processes in maize. Genes were classified into groups based on the predicted function of the encoded proteins: cell wall metabolism, cytoskeleton, general metabolism, signaling and unknown. In-situ hybridization performed for two selected genes, confirmed the spatial distribution of expression shown by Northern blots and revealed subtle differences in tissue localization. Interestingly, spatial profiles of expression for some cell wall related genes appeared to correlate with the profile of accelerating root elongation and changed appropriately under growth-inhibitory water deficit.