Phosphate Availability Alters Lateral Root Anatomy and Root Architecture of Fraxinus mandshurica Rupr. Seedlings

Plants have evolved some mechanisms to maximize the efficiency of phosphorus acquisition. Changes in root architecture are one such mechanism. When Fraxinus mandshurica Rupr. seedlings were grown under conditions of low phosphorus availability, the length of cells in the meristem zone of the lateral roots was longer, but the length of cells in the elongation and mature zones of the lateral roots was shorter,compared with seedlings grown under conditions of high phosphorus availability. The elongation rates of primary roots increased as phosphorus availability increased, but the elongation rates of the branched zones of the primary roots decreased. The number of lateral root primordia and the length of the lateral roots decreased as phosphorus availability increased. The topological index (altitude slope) decreased as phosphorus availability increased, suggesting that root architecture tended to be herringbone-like when seedlings were grown under conditions of low phosphate availability. Herringbone-like root systems exploit nutrients more efficiently, but they have higher construction costs than root systems with a branching pattem.     

Primary and Lateral Root Development of Dark- and Light-Grown Cotton Seedlings under Salinity Stress*

Primary root growth dynamics and lateral root development of dark- and light grown cotton seedlings (Gossypium hirsutum L., cv. Acala SJ-2) were studied under control and salinity stress conditions. The seedlings were grown by two methods: A) in paper-lined, vermiculite-filled beakers with the plants growing between the paper and the glass wall (Gladish and Rost, 1993), and B) in hydroponics after germination and initial growth in germination paper rolls saturated with the treatment solutions (Kent and Läuchli, 1985). After germination, daily primary root elongation rate gradually incrased to a maximum, then gradually declined to close to zero for dark-grown seedlings, or to sustained rates of about 10 mm per day for light-grown control plants. Salinity stress delayed primary root growth and reduced peak elongation rates, without changing the general primary root growth pattern. These results suggest that salinity changed the time-scale, but did not modify the normal developmental sequence. Lateral root growth was more inhibited by salinity than primary root growth. In addition, elongation of lateral roots was more inhibited by salinity than their initiation and emergence. Light exposure of the shoot favored both sustained primary root growth from 7 days after planting, and lateral root emergence and growth. Salinity effects were more severe on seedlings germinated and grown in hydroponics (method B) than on vermiculite-grown plants (method A). These results emphasize the importance of growing conditions for the NaCl-induced effects on cotton root development. In addition, the differential effects of salinity on primary and lateral roots became evident, pointing to diverse control mechanisms for the development of these root types.     

Reactive oxygen species localization in roots of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> seedlings grown under phosphate deficiency

Arabidopsis plants responding to phosphorus (P) deficiency increase lateral root formation and reduce primary root elongation. In addition the number and length of root hairs increases in response to P deficiency. Here we studied the patterns of radical oxygen species (ROS) in the roots of Arabidopsis seedlings cultured on media supplemented with high or low P concentration. We found that P availability affected ROS distribution in the apical part of roots. If plants were grown on high P medium, ROS were located in the root elongation zone and quiescent centre. At low P ROS were absent in the elongation zone, however, their synthesis was detected in the primary root meristem. The proximal part of roots was characterized by ROS production in the lateral root primordia and in elongation zones of young lateral roots irrespective of P concentration in the medium. On the other hand, plants grown at high or low P differed in the pattern of ROS distribution in older lateral roots. At high P, the elongation zone was the primary site of ROS production. At low P, ROS were not detected in the elongation zone. However, they were present in the proximal part of the lateral root meristem. These results suggest that P deficiency affects ROS distribution in distal parts of Arabidopsis roots. Under P-sufficiency ROS maximum was observed in the elongation zone, under low P, ROS were not synthesized in this segment of the root, however, they were detected in the apical root meristem.     

磷空间有效性对拟南芥根形态构型的影响

磷空间有效性显著影响拟南芥主、侧根生长。在均一的磷处理下,极度磷胁迫或过量供磷均会导致拟南芥主根变短和侧根密度降低。在分层的磷处理下,上层高磷下层低磷能明显促进主根伸长生长,提高侧根在高磷区域的密度,说明植物根系在下层低磷区感受到磷胁迫信号后,可促进上层高磷区侧根的形成和发育。     

A role for redox factors in shaping root architecture under phosphorus deficiency

The developmental response of the Arabidopsis root system to low phosphorus (P) availability involves the reduction in primary root elongation accompanied by the formation of numerous lateral roots. We studied the roles of selected redox metabolites, namely, radical oxygen species (ROS) and ascorbic acid (ASC) in the regulation of root system architecture by different P availability. Rapidly growing roots of plants grown on P-sufficient medium synthesize ROS in root elongation zone and quiescent centre. We have demonstrated that the arrest of root elongation at low P medium coincides with the disappearance of ROS from the elongation zone. P-starvation resulted in a decrease in ascorbic acid level in roots. This correlated with a decrease in cell division activity. On the other hand, feeding P-deficient plants with ASC, stimulated mitotic activity in the primary root meristem and partly reversed the inhibition of root growth imposed by low P conditions. In this paper, we discuss the idea of the involvement of redox agents in the regulation of root system architecture under low P availability.Key words: ascorbic acid, phosphate deficiency, primary root, radical oxygen species, root growth, root system architecture     

Morphometric analysis of Pinus banksiana Lamb. root anatomy during a 3-month field study

Tree roots are variable in their growth rates, alternating between periods of elongation and dormancy. This variability may have a strong influence on root anatomy. In the present study, field-grown Pinus banksiana Lamb. roots were divided into four distinct anatomical regions (i.e. white without mycorrhizae, white with mycorrhizae, condensed tannin, and cork). Changes in root growth, the proportions of the root system occupied by the various regions, and cortical plasmalemma surface area (CPSA) were determined for 6- to 9-month-old ectomycorrhizal P. banksiana seedlings during a 3-month period (August through October) in northern Ontario. The region in which the greatest change in length occurred was the condensed tannin zone, which was also the dominant contributor to root length (up to 74% of total). The roots of seedlings grown under artificial conditions had the same zones but in different proportions compared to roots in the field. A correlation was noted between increased root growth, low metacutization, and high soil water availability. The CPSA data were assumed to be a factor influencing ion uptake capacity in a positive manner. Interestingly, increases in CPSA were not directly correlated with changes in root length for field-grown seedlings. The primary contributor to CPSA in the field-grown roots was the ectomycorrhizal zone (approximately 80%). In comparison, the bulk (85%) of the CPSA in the chamber-grown roots was found in the white root region. The conditions under which the seedlings were grown strongly influenced the anatomy of their roots. Received: 27 April 1999 / Accepted: 6 December 1999     

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.     

Root growth responses to lead in young maize seedlings

This work was undertaken to follow the appearance and development of symptoms of lead toxicity in growing roots of seedlings. The effects of lead nitrate (10^-2–10⁵ M) were studied on the roots of maize (Zea mays) seedlings, cvs. Diamant and Sterling. The roots were grown on filter paper either on glass in trays or in large Petri dishes. The following characteristics of root growth were studied: seed germination, length of primary and seminal roots, number of seminal and lateral roots, length of branching zone, length of meristem and fully-elongated cells and the number of fully-elongated cells along the daily length increment. 10^-2 M lead nitrate exerted a clear toxic effect on root elongation just after radicle emergence; its influence on shoot growth was weak. However 10^-2 M Pb solution did not affect either radicle emergence itself or seminal root emergence, which can be explained by the impermeability of seed testa to lead salt. The inhibitory effect of 10^-3 M lead nitrate appeared a day later and was not as toxic: the growth of primary and seminal roots proceeded at lower rate due to a partial inhibition of cell division and cell elongation in them. 10^-3 M lead nitrate modified the root system morphology: it exerted no effect on the emergence of lateral roots and their number, but induced a more compact distribution of lateral roots along a shorter branching zone due to a reduced length of mature cells in the primary root. As a result of the more prominent inhibition of primary root growth, a shorter branching zone with more compactly located lateral roots occupied a position much closer to the root tip than in roots grown without the influence of lead.     

Adaptive Changes and Genotypic Variation for Root Architecture of Common Bean in Response to Phosphorus Deficiency

Root architectural responses to phosphorus (p) availability may be an important trait for P acquisition efficiency. In the present study, The authors examined the effects of P availability on root architectural responses of different common bean genotypes. Five common bean (Phaseolus vulgaris L.) genotypes representing different origins and ecotypic races were compared both in a specially designed paper pouch system and a stratified P buffer sand culture system with computer image analysis. The results showed that root architecture was regulated by P availability. P deficiency led to form a shallower root system, as indicated by increased relative distribution of basal root length in the upper layers and decreased the growth angle of basal roots. There was significant genetic variation in root architecture in response to P deficiency both in the paper pouch system and the stratified sand culture system. Under low P conditions some genotypes were more gravitropically sensitive to low P availability, resulting in producing a shallower root system and enhanced root exploration into the surface soil, where soil available P is more concentrated. G19833 and DOR364, which were most contrasting in P efficiency, were also very different in root architectural response to P availability. The results from this study suggest that P availability regulates root architecture and P deficiency leads to shallower root architecture in beans. The genetic potential of root architecture provides the possibility of selecting this trait for improving P acquisition efficiency in common bean.     

Detection of quantitative trait loci for seminal root traits in maize (Zea mays L.) seedlings grown under differential phosphorus levels

Suboptimal phosphorus availability is a primary constraint for terrestrial plant growth. Seminal roots play an important role in acquisition of nutrients by plant seedlings. The length and number of seminal roots may be particularly important in acquisition of immobile nutrients such as phosphorus by increasing soil exploration. The objective of this study was to identify quantitative trait loci (QTL) controlling seminal root growth in response to phosphorus stress in maize, and to characterize epistatic interactions among QTL. Seminal root length and number were evaluated in 162 recombinant inbred lines derived from a cross between B73 and Mo17 in seedlings grown in a controlled environment. B73 and Mo17 significantly differed for seminal root length under low phosphorus, but not under adequate phosphorus conditions. Seminal root length of the population grown under low phosphorus ranged from 0 to 79.2 cm with a mean of 32.3 cm; while seminal root length of plants grown under high phosphorus ranged from 0.67 to 59.0 cm with a mean of 23.4 cm. Under low phosphorus, one main-effect QTL was associated with seminal root length and three QTL with seminal root number; under high phosphorus, two QTL with seminal root length and three QTL for seminal root number. These accounted for 11, 25.4, 22.8, and 24.1% of the phenotypic variations for seminal root length and number at low phosphorus, and seminal root length and number at high phosphorus, respectively. Di-genic epistatic loci were detected for seminal root length at low phosphorus (two pairs) seminal root number at low phosphorus (eight pairs), seminal root length at high phosphorus (four pairs), and seminal root number at high phosphorus (two pairs), which accounted for 23.2, 50.6, 32.2, and 20.3% of the total variations, respectively. Seminal root traits observed here were positively yet weakly correlated with shoot biomass in the field under low phosphorus, although no coincident QTL were detected. These results suggest that epistatic interactions are important in controlling genotypic variation associated with seedling seminal root traits.     

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.     

Phosphate availability alters architecture and causes changes in hormone sensitivity in the Arabidopsis root system

The postembryonic developmental program of the plant root system is plastic and allows changes in root architecture to adapt to environmental conditions such as water and nutrient availability. Among essential nutrients, phosphorus (P) often limits plant productivity because of its low mobility in soil. Therefore, the architecture of the root system may determine the capacity of the plant to acquire this nutrient. We studied the effect of P availability on the development of the root system in Arabidopsis. We found that at P-limiting conditions (<50 microM), the Arabidopsis root system undergoes major architectural changes in terms of lateral root number, lateral root density, and primary root length. Treatment with auxins and auxin antagonists indicate that these changes are related to an increase in auxin sensitivity in the roots of P-deprived Arabidopsis seedlings. It was also found that the axr1-3, axr2-1, and axr4-1 Arabidopsis mutants have normal responses to low P availability conditions, whereas the iaa28-1 mutant shows resistance to the stimulatory effects of low P on root hair and lateral root formation. Analysis of ethylene signaling mutants and treatments with 1-aminocyclopropane-1-carboxylic acid showed that ethylene does not promote lateral root formation under P deprivation. These results suggest that in Arabidopsis, auxin sensitivity may play a fundamental role in the modifications of root architecture by P availability.     

Inhibition of ammonium assimilation restores elongation of seminal rice roots repressed by high levels of exogenous ammonium

Elongation of seminal and lateral roots of rice seedlings was markedly inhibited by high ammonium levels in growth medium. However, high exogenous nitrate concentrations had little inhibitory effect on root growth. The objective of this study was to elucidate the relationship between inhibition of rice root growth induced by high ammonium conditions and ammonium assimilation in the seedlings. Activity of glutamine synthetase (GS) was kept at a low level in the seminal roots of the seedlings grown under high nitrate levels. In contrast, high ammonium levels significantly enhanced the GS activity in the roots, so that Gln abundantly accumulated in the shoots. These results indicate that ammonium assimilation may be activated in the seminal roots under high ammonium conditions. Application of methionine sulfoximine (MSO), an inhibitor of GS, relieved the repression of the seminal root elongation induced by high ammonium concentrations. However, the elongation of lateral roots remained inhibited even under the same condition. Furthermore, MSO drastically increased ammonium level and remarkably decreased Gln level in the shoots grown under high ammonium conditions. These results show that, for rice seedlings, an assimilatory product of ammonium, and not ammonium itself, may serve as an endogenous indicator of the nitrogen status involved in the inhibition of seminal root elongation induced by high levels of exogenous ammonium.     

Root growth, developmental changes in the apex, and hydraulic conductivity for Opuntia ficus-indica during drought

Developmental changes in the root apex and accompanying changes in lateral root growth and root hydraulic conductivity were examined for Opuntia ficus-indica (L.) Miller during rapid drying, as occurs for roots near the soil surface, and more gradual drying, as occurs in deeper soil layers. During 7 d of rapid drying (in containers with a 3-cm depth of vermiculite), the rate of root growth decreased sharply and most root apices died; such a determinate pattern of root growth was not due to meristem exhaustion but rather to meristem mortality after 3 d of drying. The length of the meristem, the duration of the cell division cycle, and the length of the elongation zone were unchanged during rapid drying. During 14 d of gradual drying (in containers with a 6-cm depth of vermiculite), root mortality was relatively low; the length of the elongation zone decreased by 70%, the number of meristematic cells decreased 30%, and the duration of the cell cycle increased by 36%. Root hydraulic conductivity ( L _P) decreased to one half during both drying treatments; L _P was restored by 2 d of rewetting owing to the emergence of lateral roots following rapid drying and to renewed apical elongation following gradual drying. Thus, in response to drought, the apical meristems of roots of O. ficus-indica near the surface die, whereas deeper in the substrate cell division and elongation in root apices continue. Water uptake in response to rainfall in the field can be enhanced by lateral root proliferation near the soil surface and additionally by resumption of apical growth for deeper roots.     

Regulation of root hair density by phosphorus availability in Arabidopsis thaliana

We characterized the response of root hair density to phosphorus (P) availability in Arabidopsis thaliana. Arabidopsis plants were grown aseptically in growth media with varied phosphorus concentrations, ranging from 1 mmol m⁻³ to 2000 mmol m⁻³ phosphorus. Root hair density (number of root hairs per mm of root length) was analysed starting at 7 d of growth. Root hair density was highly regulated by phosphorus availability, increasing significantly in roots exposed to low-phosphorus availability. The initial root hairs produced by the radicle were not sensitive to phosphorus availability, but began to respond after 9 d of growth. Root hair density was about five times greater in low phosphorus (1 mmol m⁻³) than in high phosphorus (1000 mmol m⁻³) media. Root hair density decreased logarithmically in response to increasing phosphorus concentrations within that range. Root hair density also increased in response to deficiencies of several other nutrients, but not as strongly as to low phosphorus. Indoleacetic acid (IAA), the auxin transport inhibitor 2-(p-chlorophenoxy)-2-methylpropionic acid (CMPA), the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC), and the ethylene synthesis inhibitor amino-oxyacetic acid (AOA) all increased root hair density under high phosphorus but had very little effect under low phosphorus. Low phosphorus significantly changed root anatomy, causing a 9% increase in root diameter, a 31% decrease in the cross-sectional area of individual trichoblasts, a 40% decrease in the cross-sectional area of individual atrichoblasts, and 45% more cortical cells in cross-section. The larger number of cortical cells and smaller epidermal cell size in low phosphorus roots increased the number of trichoblast files from eight to 12. Two-thirds of increased root hair density in low phosphorus roots was caused by increased likelihood of trichoblasts to form hairs, and 33% of the increase was accounted for by changes in low phosphorus root anatomy resulting in an increased number of trichoblast files. These results show that phosphorus availability can fundamentally alter root anatomy, leading to changes in root hair density, which are presumably important for phosphorus acquisition.     

Phosphate starvation induces a determinate developmental program in the roots of Arabidopsis thaliana

When growing under limiting phosphate (P) conditions, Arabidopsis thaliana plants show dramatic changes in root architecture, including a reduction in primary root length, increased formation of lateral roots and greater formation of root hairs. Here we report that primary root growth inhibition by low P is caused by a shift from an indeterminate to a determinate developmental program. In the primary root, the low P-induced determinate growth program initiates with a reduction of cell elongation followed by the progressive loss of meristematic cells. At later stages, cell proliferation ceases and cell differentiation takes place at the former cell elongation and meristematic regions of the primary root. In low P, not only the primary but also almost all mature lateral roots enter the determinate developmental program. Kinetic studies of expression of the cell cycle marker CycB1;1:uidA and the quiescent center (QC) identity marker QC46:GUS showed that in low P conditions, reduction in proliferation in the primary root was preceded by alterations in the QC. These results suggest that in Arabidopsis, P limitation can induce a determinate root developmental program that plays an important role in altering root system architecture and that the QC could act as a sensor of environmental signals.     

低磷供应对拟南芥根系构型的影响

在人工气候箱中,采用Johnson培养基对拟南芥在低磷供应条件下根系构型的变化进行了研究,结果表明：拟南芥在磷饥饿诱导下,主根缩短,侧根密度、根毛的数量和长度显著增加,并且,根尖到第一侧根和第一根毛的距离也大大缩短。这些改变增加了根系比表面积,并且使得根系分布更加靠近土壤表层,有利于提高植物吸收土壤中有机磷的效率。低磷胁迫还导致拟南芥根系分生组织区细胞形状变异,柱细胞数量减少;主根生长和细胞伸长的动力学分析显示,磷饥饿促使拟南芥主根生长变缓,细胞长度随磷饥饿程度的加深迅速缩小。CycB1;1:GUS染色分析结果表明,低磷破坏拟南芥根系分生组织细胞分裂能力,这些结果说明磷胁迫同时抑制了细胞的伸长和分裂,从而引起拟南芥主根的缩短。     

Ethylene: a regulator of root architectural responses to soil phosphorus availability

The involvement of ethylene in root architectural responses to phosphorus availability was investigated in common bean ( Phaseolus vulgaris L.) plants grown with sufficient and deficient phosphorus. Although phosphorus deficiency reduced root mass and lateral root number, main root length was unchanged by phosphorus treatment. This resulted in decreased lateral root density in phosphorus-deficient plants. The possible involvement of ethylene in growth responses to phosphorus deficiency was investigated by inhibiting endogenous ethylene production with amino-ethoxyvinylglycine (AVG) and aerating the root system with various concentrations of ethylene. Phosphorus deficiency doubled the root-to-shoot ratio, an effect which was suppressed by AVG and partially restored by exogenous ethylene. AVG increased lateral root density in phosphorus- deficient plants but reduced it in phosphorus-sufficient plants. These responses could be reversed by exogenous ethylene, suggesting ethylene involvement in the regulation of main root extension and lateral root spacing. Phosphorus-deficient roots produced twice as much ethylene per g dry matter as phosphorus-sufficient roots. Enhanced ethylene production and altered ethylene sensitivity in phosphorus-deficient plants may be responsible for root responses to phosphorus deficiency.     

Growth and morphology of eucalypt seedling-roots, in relation to soil strength arising from compaction

Information on the response of root growth and morphology to soil strength is useful for testing suitability of existing and new tillage methods and/or for selecting plants suitable for a specific site with or without tillage. Although there is extensive published information on the root growth-soil strength relationships for annual agricultural plants, such information is scarce for woody, perennial tree species. The purpose of this study is to examine growth and morphology of the root systems of 17-day-old eucalypt seedlings with respect to variation in soil strength. Soil strength in this study was varied by compaction of a well-aggregated clay soil to bulk densities of 0.7–1.0 Mg m^-3 whilst maintaining adequate water availability and aeration for plant growth. Lengths and tip-diameters of primary and lateral roots were measured on the excavated root systems of seedlings.With increase in bulk density and also soil strength (expressed as penetrometer resistance), total length of primary and lateral roots decreased. There were 71 and 31% reduction in the lengths of primary and lateral roots respectively with an increase in penetrometer resistance from 0.4 to 4.2 MPa. This indicated primary roots to be more sensitive to high soil strength than the lateral roots. Average length of lateral roots and diameters of both primary and lateral root tips increased with an increase in soil strength as well. There was greater abundance of lateral roots (no. of lateral roots per unit length of primary root) and root hairs with increased soil strength. The observed root behaviour to variable soil strength is discussed in the context of compensatory growth of roots and overall growth of plants.