A comprehensive analysis of root morphological changes and nitrogen allocation in maize in response to low nitrogen stress

The plasticity of root architecture is crucial for plants to acclimate to unfavourable environments including low nitrogen (LN) stress. How maize roots coordinate the growth of axile roots and lateral roots (LRs), as well as longitudinal and radial cell behaviours in response to LN stress, remains unclear. Maize plants were cultivated hydroponically under control (4 mm nitrate) and LN (40 μm ) conditions. Temporal and spatial samples were taken to analyse changes in the morphology, anatomical structure and carbon/nitrogen (C/N) ratio in the axile root and LRs. LN stress increased axile root elongation, reduced the number of crown roots and decreased LR density and length. LN stress extended cell elongation zones and increased the mature cell length in the roots. LN stress reduced the cell diameter and total area of vessels and increased the amount of aerenchyma, but the number of cell layers in the crown root cortex was unchanged. The C/N ratio was higher in the axile roots than in the LRs. Maize roots acclimate to LN stress by optimizing the anatomical structure and N allocation. As a result, axile root elongation is favoured to efficiently find available N in the soil.     

硝酸盐供应对玉米侧根生长的影响

以两个玉米(Zea mays L.)自交系478和Wu312为研究材料,采用琼脂培养方法,研究不同浓度NO-3对侧根生长的影响.结果表明,在外部浓度0.01～1.0mmol/L范围内,NO-3供应能显著增加侧根的长度及根生物量.但当NO-3供应超过1.0 mmo1/L后,侧根长度开始下降.当NO-3供应分别在超过5.0(Wu312)与10(478)mmol/L后,侧根密度显著下降.在10 mmol/LNO-3下,Wu312的侧根生长几乎完全被抑制.而478在20 mmol/L时,侧根密度仍可达到其最大值的30%(主根)～50%(胚根).植株地上部全氮及硝酸盐含量随NO-3供应的增加而升高,二者与侧根长度、侧根密度及冠根比的数学函数关系相同.     

Genetic Analysis of Maize Root Characteristics in Response to Low Nitrogen Stress

Under low-input cropping systems, nitrogen (N) can be a limiting factor in plant growth and yield. Identifying genotypes that are more efficient at capturing limited N resources and the traits and mechanisms responsible for this ability is important. Root trait has a substantial influence on N acquisition from soils. Nevertheless, inconsistencies still exist as to the effect of low N on root length and its architecture in terms of lateral and axial roots. For maize, a crop utilizing heterosis, little is known about the relationship between parents and their crosses in the response of root architecture to N availability. Here 7 inbred maize lines and 21 of their crosses created by diallel mating were used to study the effect of N stress on root morphology as well as the relationship between the inbreds and their crosses. With large genotypic differences, low N generally suppresses shoot growth and increases the root to shoot ratio with or without increasing root biomass in maize. Maize plants responded to N deficiency by increasing total root length and altering root architecture by increasing the elongation of individual axial roots and enhancing lateral root growth, but with a reduction in the number of axial roots. Here, the inbreds showed weaker responses in root biomass and other root parameters than their crosses. Heterosis of root traits was significant at both N levels and was attributed to both the general combining ability (GCA) and special combining ability (SCA). Low N had substantial affects on the pattern of heterosis, GCA and SCA affects on root traits for each of the crosses suggesting that selection under N stress is necessary in generating low N-tolerant maize genotypes.     

常绿阔叶林外生和丛枝菌根树种细根形态和构型性状对氮添加的可塑性响应

以中亚热带常绿阔叶林外生菌根树种罗浮栲和丛枝菌根树种木荷为研究对象,采用根袋法进行野外原位氮添加试验,研究了细根形态性状(比根长、比表面积、组织密度、平均根直径)和构型性状(分枝数、分枝比、根长增长速率、根尖密度、分枝密度),分析不同菌根树种细根形态和构型性状对氮沉降的响应.结果表明:随序级增加,外生和丛枝菌根树种细根...     

Lateral root initiation in Arabidopsis: developmental window, spatial patterning, density and predictability

BACKGROUND AND AIMS: The basic regulatory mechanisms that control lateral root (LR) initiation are still poorly understood. An attempt is made to characterize the pattern and timing of LR initiation, to define a developmental window in which LR initiation takes place and to address the question of whether LR initiation is predictable. METHODS: The spatial patterning of LRs and LR primordia (LRPs) on cleared root preparations were characterized. New measures of LR and LRP densities (number of LRs and/or LRPs divided by the length of the root portions where they are present) were introduced and illustrate the shortcomings of the more customarily used measure through a comparative analysis of the mutant aux1-7. The enhancer trap line J0121 was used to monitor LR initiation in time-lapse experiments and a plasmolysis-based method was developed to determine the number of pericycle cells between successive LRPs. KEY RESULTS: LRP initiation occurred strictly acropetally and no de novo initiation events were found between already developed LRs or LRPs. However, LRPs did not become LRs in a similar pattern. The longitudinal spacing of lateral organs was variable and the distance between lateral organs was proportional to the number of cells and the time between initiations of successive LRPs. There was a strong tendency towards alternation in LR initiation between the two pericycle cell files adjacent to the protoxylem poles. LR density increased with time due to the emergence of slowly developing LRPs and appears to be unique for individual Arabidopsis accessions. CONCLUSIONS: In Arabidopsis there is a narrow developmental window for LR initiation, and no specific cell-count or distance-measuring mechanisms have been found that determine the site of successive initiation events. Nevertheless, the branching density and lateral organ density (density of LRs and LRPs) are accession-specific, and based on the latter density the average distance between successive LRs can be predicted.     

System analysis of microRNAs in the development and aluminium stress responses of the maize root system

MicroRNAs (miRNAs) are a class of regulatory small RNAs (sRNAs) that down‐regulate target genes through mRNA cleavage or translational inhibition. miRNA is known to play an important role in the root development and environmental responses in both the Arabidopsis and rice. However, little information is available to form a complete view of miRNAs in the development of the maize root system and Al stress responses in maize. Four sRNA libraries were generated and sequenced from the early developmental stage of primary roots (PRY), the later developmental stage of maize primary roots (PRO), seminal roots (SR) and crown roots (CR). Through integrative analysis, we identified 278 miRNAs (246 conserved and 32 novel ones) and found that the expression patterns of miRNAs differed dramatically in different maize roots. The potential targets of the identified conserved and novel miRNAs were also predicted. In addition, our data showed that CR is more resistant to Al stress compared with PR and SR, and the differentially expressed miRNAs are likely to play significant roles in different roots in response to environmental stress such as Al stress. Here, we demonstrate that the expression patterns of miRNAs are highly diversified in different maize roots. The differentially expressed miRNAs are correlated with both the development and environmental responses in the maize root. This study not only improves our knowledge about the roles of miRNAs in maize root development but also reveals the potential role of miRNAs in the environmental responses of different maize roots.     

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.     

杨树人工林细根数量和形态特征的季节动态及代际差异

采集欧美杨107Ⅰ代和Ⅱ代人工林细根样品,分析杨树不同根序细根数量特征(根长度、表面积和生物量)和形态特征(比根长、根长密度、根组织密度)对季节波动的响应及其代际差异.结果表明： 杨树各根序细根数量特征(根长度、表面积和生物量)均呈明显的季节变化,且具有明显的根序差异性.低级根序细根数量特征季节差异显著,细根生物量在生长季显著增加而生长季后显著下降.高级根序细根比根长季节波动显著,而根长密度和根组织密度等形态特征波动较小.连作导致人工林杨树1～2级细根长度、生物量、比根长和根长密度在生长季显著增大.1级细根数量特征与土壤温湿度呈显著正相关,与土壤有机质和速效氮含量呈显著负相关;而2级细根数量特征仅与土壤养分显著相关.杨树人工林细根特征的季节动态及代际差异体现了杨树对细根的碳投入变化,因连作引发的土壤养分匮乏可能引发植株对根系的碳投入增加,这种碳分配格局与人工林地上部分生产力形成密切相关.     

Root and leaf functional trait relations in Poaceae species: implications of differing resource-acquisition strategies

Aims Root systems play an essential role in grassland functioning in both acquisition and storage of resources. Nevertheless, root functional traits have not received as much attention as those measured on above-ground organs, and little is known about their relations. Our objectives were to test whether morphological and root system traits allowed identification of grass species' functional strategies and to determine whether a relation exists between above- and below-ground traits.Methods Functional traits of root tissues (specific root length, diameter, tissue density and nitrogen concentration), whole root systems (root mass, root length density, root mass percentage below a depth of 20cm and fine root %) and two major leaf traits (specific leaf area and leaf dry matter content) were determined under field conditions and their relations were analysed in eleven perennial temperate Poaceae species.Important findings Canonical correspondence analysis along Axis 1 revealed a gradient of species, from those with deep, dense and coarse root systems with a large root mass to those with shallow root systems, thin roots and high specific root length; this suggests strong correlations among root traits. Correlations between specific root length and specific leaf area reveal two groups of species, which probably indicates different drought-tolerance capacities. Root trait syndromes enable ranking grasses along a gradient from conservative-strategy species (from stressful habitats), which display a deep and coarse root system, to acquisitive species (from rich and moist meadows), which display a shallow and thin root system. Although both types display similar above-ground strategies, drought-tolerant species have lower specific root lengths than drought-sensitive species, revealing more conservative root strategies.     

Root Growth of the Annual Tillering Grass Panicum miliaceum in Heterogeneous Nutrient Environments (in English)

To study growth responses of the roots of Panicum miliaceum L. to heterogeneous supply of nutrients. The authors analyzed the effects of the nutrient levels in both original patches (O) and destination patches (D) on the root growth of P. miliaceum when its roots were allowed to extend from original patch into destination patch. When the nutrient levels in the original patches were low, coarse root biomass ratio (coarse root biomass in the D/total coarse root biomass), coarse root length ratio (coarse root length in the D/total coarse root length), coarse root surface area ratio (coarse root surface area in the D/total coarse root surface area) and fine root length ratio (fine root length in the D/total fine root length) were greater in the destination patches with lower nutrient levels than in the destination patches with higher nutrient levels, while fine root length, fine root length density, fine root surface index, and fine root surface area density were smaller in the former than in the latter. When the nutrient levels in the original patches were high, fine root length, fine root length density, fine root surface area index and fine root surface density were greater in the destination patches with lower nutrient levels than in the destination patches with higher nutrient levels, coarse roots did not respond to the nutrient levels in the destination patches significantly. When the roots extended from the original patches with the same nutrient level into the destination patches with contrasting nutrient levels, fine root biomass and its percentage allocation did not respond to the nutrient levels in the destination patches significantly, whereas both root length and root surface area did. This indicates that the fine roots of P. miliaceum responded to difference in nutrient supply by plasticity in their length and surface area, rather than in their root biomass.     

异质养分环境中一年生分蘖草本黍根系的生长特征

为揭示黍（Panicum miliaceum L.)根系对异质养分环境的生长反应,作研究了黍根系从起始斑块向目标斑块水平生长时,时始斑块和目标斑块养分水平根生长的影响,就低养分起始珏块而言,粗根生物量,粗根长度,粗根表面积和细极长度在高养分目标斑块中的分配比例均小于其在低养分目标斑块中的分配比例,而细根长度及其密度,细根表面积指及其密度的变化恰好相反,就高养分起始斑块而言,高养分目标斑块的细根长度,细根长度密度,细根表面积指数和细根表面积密均不于低养分目标斑块,而粗根对目标斑块中养分状的反应不明显。当黍根系从桢的起始斑块进入不同的目标斑块后,目标斑块的养分状况对细根生物量及其分配无影响,而显影响细根长度和表现积,这指示细根是通过长度和表面积可塑性而不是生物量变化响应目标斑块中的养分差异。     

黄土丘陵区植物叶片与细根功能性状关系及其变化

通过植物叶片功能性状(比叶面积、叶组织密度、叶氮含量)和细根功能性状(比根长、根组织密度、根氮含量)间的相互关系,分析植物对环境的适应途径;然后根据性状间的差异进行了层次聚类,将物种划分为3大功能型,并分析了不同功能型对环境的适应策略.结果表明:黄土丘陵区延河流域149种植物的叶氮含量与比叶面积和根氮含量正相关、与叶组织密度负相关,比根长与根组织密度负相关,除了根氮含量,其余根性状与叶性状不相关.此外,功能性状间关系变化和适应策略在不同功能型之间也存在差异.功能型1的植物具有最强的耐旱力和防御力;功能型3的植物具有最强的养分维持能力用以对抗营养贫瘠的环境;功能型2的植物居中,生长速率最高,具有较强的竞争力、分布最广;根据C-S-R理论,功能型1和3属于“胁迫忍耐型”策略(S策略),功能型2则属于“竞争型”(C)和“干扰型”(R)策略的综合.研究结果为黄土丘陵区植被恢复规划及物种配置等提供依据.     

Cadmium stress suppresses lateral root formation by interfering with the root clock

A biological clock activated by oscillating signals, known as root clock, has been linked to lateral root (LR) formation and is essential for regular LR spacing along the primary root. However, it remains unclear how this internal mechanism is influenced by environmental factors known to affect the LR pattern. Here, we report that excessive cadmium (Cd) inhibits LR formation by disrupting the lateral root cap (LRC)‐programmed cell death (PCD)‐regulated root clock. Cd restricts the frequency of the oscillating signal rather than its amplitude. This could be attributed to the inhibition on meristematic activity by Cd, which resulted in decreased LRC cell number and LRC‐PCD frequency. Genetic evidence further showed that LRC cell number is positively correlated with root resistance to Cd. Our study reveals root cap dynamics as a novel mechanism mediating root responses to Cd, providing insight into the signalling pathways of the root clock responding to environmental cues.     

Phenotypic plasticity of <Emphasis Type="Italic">Chenopodium murale</Emphasis> across contrasting habitat conditions in peri-urban areas in Indian dry tropics: Is it indicative of its invasiveness?

Phenotypic plasticity is an important plant trait associated with invasiveness of alien plants that reflects its ability to occupy a wide range of environments. We investigated the phenotypic response of Chenopodium murale to resource variability and ontogeny. Its plant-level and leaf-level traits were studied at high-resource (HR) and low-resource (LR) sites in peri-urban areas in Indian dry tropics. Plants at LR had significantly higher root length, root/shoot biomass ratio, stem mass and root mass fractions. Plants at HR had higher shoot length, basal diameter, leaf mass fraction and leaf area ratio. Leaf-level traits like leaf area and chlorophyll a were also higher here. Mean plasticity indices for plant- and leaf-level traits were higher at HR. With increasing total plant biomass, there was significant increase in the biomass of leaf, stem, root, and reproductive parts, and root and shoot lengths, whereas root/shoot length ratio, their biomass ratio, and leaf and root mass fractions declined significantly. Allocation to roots and leaves significantly decreased with increasing plant size at both sites. But, at any size, allocation to roots was greater at LR, indicative of optimization of capture of soil nutrients, whereas leaf allocation was higher at HR. Consistently increasing stem allocation equaled leaf allocation at comparatively higher shoot lengths at HR. Reproductive biomass comprised 10–12% of the plant’s total biomass. In conclusion, the success of alien weed C. murale across environmentally diverse habitat conditions in Indian dry tropics can be attributed to its high phenotypic plasticity, resource utilization capability in low-resource habitats and higher reproductive potential. These characteristics suggest that it will continue to be an aggressive invader.     

Root hairs are the most important root trait for rhizosheath formation of barley (Hordeum vulgare), maize (Zea mays) and Lotus japonicus (Gifu)

Background and AimsRhizosheaths are defined as the soil adhering to the root system after it is extracted from the ground. Root hairs and mucilage (root exudates) are key root traits involved in rhizosheath formation, but to better understand the mechanisms involved their relative contributions should be distinguished.MethodsThe ability of three species [barley (Hordeum vulgare), maize (Zea mays) and Lotus japonicus (Gifu)] to form a rhizosheath in a sandy loam soil was compared with that of their root-hairless mutants [bald root barley (brb), maize root hairless 3 (rth3) and root hairless 1 (Ljrhl1)]. Root hair traits (length and density) of wild-type (WT) barley and maize were compared along with exudate adhesiveness of both barley and maize genotypes. Furthermore, root hair traits and exudate adhesiveness from different root types (axile versus lateral) were compared within the cereal species.Key ResultsPer unit root length, rhizosheath size diminished in the order of barley > L. japonicus > maize in WT plants. Root hairs significantly increased rhizosheath formation of all species (3.9-, 3.2- and 1.8-fold for barley, L. japonicus and maize, respectively) but there was no consistent genotypic effect on exudate adhesiveness in the cereals. While brb exudates were more and rth3 exudates were less adhesive than their respective WTs, maize rth3 bound more soil than barley brb. Although both maize genotypes produced significantly more adhesive exudate than the barley genotypes, root hair development of WT barley was more extensive than that of WT maize. Thus, the greater density of longer root hairs in WT barley bound more soil than WT maize. Root type did not seem to affect rhizosheath formation, unless these types differed in root length.ConclusionsWhen root hairs were present, greater root hair development better facilitated rhizosheath formation than root exudate adhesiveness. However, when root hairs were absent root exudate adhesiveness was a more dominant trait.     

Investment in Fine Roots and Arbuscular Mycorrhizal Fungi Decrease During Succession in Three Brazilian Ecosystems

The functional groups of plants that characterize different phases of succession are expected to show differences in root distribution, fine‐root traits and degrees of association with arbuscular mycorrhizal (AM) fungi. The relationship involving fine‐root traits and AM fungi that regulate the nutrient acquisition potential among different plant functional groups are still not well understood. We assessed fine‐root morphology, AM fungal variables and soil fertility in grassland, secondary forest and mature forest in Atlantic, Araucaria and Pantanal ecosystems in Brazil. Soil cores were collected at 0–10 and 10–20 cm depths. Fine roots were extracted from soil by sieving and root morphological traits and AM colonization were determined. The AM spores were extracted from soil and counted. In all ecosystems, soil fertility, fine‐root mass and root diameter increased with the succession, while root length, specific root length, root‐hair length, root‐hair incidence, AM colonization and AM spore density decreased. These results suggest that plant species from early stages of tropical succession with inherent rapid growth invest in fine roots and maintain a high degree of AM colonization in order to increase the capacity for nutrient acquisition. Conversely, fine root morphological characteristics and low degree of AM colonization exhibited by plants of the later stages of succession lead toward a low nutrient uptake capacity that combine with their typical low growth rates. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

The impact of limited soil moisture and waterlogging stress conditions on morphological and anatomical root traits in maize (Zea mays L.) hybrids of different drought tolerance

Effects of soil drought or waterlogging on the morphological traits of the root system and internal root anatomy were studied in maize hybrids of different drought tolerance. The investigations comprised quantitative and qualitative analyses of a developed plant root system through determining the number, length and dry matter of the particular components of the root system and some traits of the anatomical structure of the seminal root. Obtained results have demonstrated a relatively broad variation in the habit of the root system. This mainly refers, to the number, length and dry matter of lateral roots, developed by seminal root, seminal adventitious and nodal roots as well as to some anatomical properties of the stele, cortex and metaxylem elements. Plants grown under waterlogging or drought conditions showed a smaller number and less dry matter of lateral branching than plants grown in control conditions. The harmful effect of waterlogging conditions on the growth of roots was greater when compared with that of plants exposed to drought. In the measurements of the root morphological traits, the effect of soil drought on the internal root anatomical characteristic was weaker than the effect of soil waterlogging. The observed effects of both treatments were more distinct in a drought sensitive hybrid Pioneer D than in drought resistant Pioneer C one. The drought resistant hybrid Pioneer C distinguished by a more extensive rooting and by smaller alterations in the root morphology caused by the stress conditions than drought sensitive hybrid Pioneer D one. Also the differences between the resistant and the sensitive maize hybrids were apparent for examined root anatomical traits. Results confirm that the hybrid Pioneer D of a high drought susceptibility was found to be also more sensitive to periodieal soil water excess. A more efficient water use and a lower shoot to root (S:R) ratio were found to be major reasons for a higher stress resistance of the hybrid Pioneer C. The reasons for a different response of the examined hybrids to the conditions of drought or waterlogging may be a more economical water balance and more favourable relations between the shoot and root dimensions in the drought resistant genotype. The observed modifications of the internal root structure caused by water deficit in plant tissues may partly influence on water conductivity and transport within roots. The results suggest that the morphological and anatomical traits of the maize root system may be used in practice as direct or indirect selection criteria in maize breeding.     

硝态氮（NO3^—）对水稻侧根生长及其氮吸收的影响

侧根是植物吸收利用土壤养分的重要器官 ,其生长发育受内部遗传因子和外部环境矿质养分的影响。通过琼脂分层培养发现 :局部供应NO-3 可以诱导水稻 (OryzasativaL .)主根或不定根上侧根的生长。为研究旱种条件下NO-3 对水稻侧根发育及其N吸收的影响 ,设置了 3个蛭石培养实验 :分根处理、全株缺N、全株供N处理。分根处理 (一半根系供应 3mmol/LKNO3,另一半根系供应 3mmol/LKCl)结果表明 :局部供应NO-3 能够促进水稻侧根生长。而在全株处理下 ,N饥饿诱导了侧根的伸长。水稻根系对NO-3 的这两种反应都存在着显著的基因型差异。同时对地上部N浓度、可溶性总糖含量及N含量分析表明 ,这些生理指标在分根处理与全株加N处理中的差异均不显著 ,表明分根处理也能基本满足植株正常生长对N的需求。在分根处理中 ,水稻的N含量与分根处理中供N一侧的平均侧根长度存在显著正相关 ,这表明在养分不均一的介质中 ,侧根长度对水稻N素吸收具有十分重要的作用。而在N素充足的条件下 ,两者之间的相关性并不显著 ,这暗示在养分充足的环境下 ,侧根长度可能并不是决定根系吸收N素的主要因素     

Scaling of root length density of maize in the field profile

Root length density is an important parameter in crop growth simulation and in evaluating consequences of root pattern on crop water and nutrient uptake. In this study, a scaling model was presented for estimating the profile distribution of root length density of maize (Zea mays L.). The model inputs are root length data of a reference profile and bulk densities of soil layers, as well as root length data in the first soil layer of a field profile to be investigated. Using the root length data of 10 soil profiles investigated over 2 years, the model was examined. The results show that the proposed scaling approach is effective in estimating the root length density of each layer of soil in the field profile. The relative root mean square error (RRMSE) of the developed scaling model was 25.28%, while that of the traditional exponential model was 39.53%. The scaling approach would facilitate determination of heterogeneous distributions of root length densities in the field.