生长素通过MAPK介导的超长链脂肪酸合成调控侧根发育

促分裂原活化蛋白激酶(MAPK)信号级联通路是真核生物中高度保守的重要信号系统,通过激酶逐级磷酸化传递并放大上游信号,进而调控细胞反应。MAPK信号通路不仅介导植物响应环境变化,而且在调节植物生长发育过程中发挥重要作用。近期,山东大学丁兆军课题组研究发现,植物重要激素生长素能够通过激活MPK14调控下游ERF13的磷酸...     

Different Ways of Formation of Root System in Plants,Maize and Flax Seedlings Taken as Examples

Comparison of the appearance and development of lateral roots in the flax and maize seedlings has shown the way of root branching in the flax, as distinct from that in most plants. Some primordia in the flax main root did not develop immediately into lateral roots, but remained quiescent, which determines different reactions of the maize and flax root systems to experimental influences. Decapitation of the main root in the maize did not leads to a significant increase in the number of lateral roots, while in the flax, their number noticeably increased due to the development of previously quiescent primordia into lateral roots. The treatment with synthetic auxin did not induce the formation of additional primordia and lateral roots in the maize roots. In the flax, the number of primordia increased significantly and that of lateral roots increased to a somewhat lesser extent. Apparently, the development of a primordium into a lateral root proceeds in two stages and they have different regulation.     

Auxin Induced Lateral Root Formation in Chicory

The supply of auxins [2,4-dichlorophenoxy acetic acid (2,4D),indole-3 acetic acid (1AA) and -naphthaleneacetic acid (NAA)]to excised chicory roots induced the formation of lateral rootmeristems mainly located close to the pre-existing apical rootmeristem. Lateral root growth induced in non-excised roots requiredhigher auxin concentrations. Inhibition of root elongation andconcomittant enlargement of the apices was also observed. SupplyingIAA induced the formation of lateral meristems earlier thanNAA, but subsequently favoured root elongation. Conversely,in the presence of 2,4D, reactivation of pericycle cells wasvery intense, but conversion of primordia to laterals was inhibited.Regardless of the auxin used, the responsive area in which lateralmeristems appeared was located a maximum of 4 mm away from theapical meristem. This region remained devoid of any lateralroot formation under control conditions. Pericycle cells oppositethe xylem poles in the diarch stele regained meristematic activityand divided transversally, giving rise to shorter cells. Thesecells subsequently divided periclinally, forming pairs of cellson the same transverse level. The root primordium extruded throughcortical cells and was surrounded by a lacuna formed to thedetriment of cortical cells.Copyright 1998 Annals of BotanyCompany Auxins,Cichorium intybus, chicory, lateral root, root elongation.     

Different methods of forming root systems in plants using maize and flax seedlings as examples

Comparison of the appearance and development of lateral roots in the flax and maize seedlings has shown the way of root branching in the flax, as distinct from that in most plants. Some primordia in the flax main root did not develop immediately into lateral roots, but remained quiescent, which determines different reactions of the maize and flax root systems to experimental influences. Decapitation of the main root in the maize did not leads to a significant increase in the number of lateral roots, while in the flax, their number noticeably increased due to the development of previously quiescent primordia into lateral roots. The treatment with synthetic auxin did not induce the formation of additional primordia and lateral roots in the maize roots. In the flax, the number of primordia increased significantly and that of lateral roots increased to a somewhat lesser extent. Apparently, the development of a primordium into a lateral root proceeds in two stages and they have different regulation.     

Some Effects of Indol-3-yl Acetic Acid on Lateral Root Development in Attached and Excised Roots of Pisum sativum L.

The effect of continuous exposure to indol-3-yl acetic acid(IAA) on primordium initiation and their subsequent emergenceas lateral roots was determined in excised and attached rootsof Pisum sativum. IAA was found to stimulate the number of primordiainitiated per centimetre of attached or excised primary. Similarly,lateral emergence in terms of the number produced per centimetreof primary was promoted in the presence of IAA. This stimulationof lateral emergence even took place in excised roots whichwere 1 cm in length at the onset of culture and which neverproduced secondary roots over a 6-d culture period when grownin the absence of auxin. These effects of IAA on lateral rootdevelopment have been considered in relation to the concurrentchanges which take place in proliferative activity in the apicalmeristem of the primary root during exposure to auxin. Pisum sativum, garden pea, anlage, primordium, emerged lateral, cell proliferation, indol-3-yl acetic acid     

Differential effects of sucrose and auxin on localized phosphate deficiency-induced modulation of different traits of root system architecture in Arabidopsis

Phosphorus, one of the essential elements for plants, is often a limiting nutrient in soils. Low phosphate (Pi) availability induces sugar-dependent systemic expression of genes and modulates the root system architecture (RSA). Here, we present the differential effects of sucrose (Suc) and auxin on the Pi deficiency responses of the primary and lateral roots of Arabidopsis (Arabidopsis thaliana). Inhibition of primary root growth and loss of meristematic activity were evident in seedlings grown under Pi deficiency with or without Suc. Although auxin supplementation also inhibited primary root growth, loss of meristematic activity was observed specifically under Pi deficiency with or without Suc. The results suggested that Suc and auxin do not influence the mechanism involved in localized Pi sensing that regulates growth of the primary root and therefore delineates it from sugar-dependent systemic Pi starvation responses. However, the interaction between Pi and Suc was evident on the development of the lateral roots and root hairs in the seedlings grown under varying levels of Pi and Suc. Although the Pi+ Suc- condition suppressed lateral root development, induction of few laterals under the Pi- Suc- condition point to increased sensitivity of the roots to auxin during Pi deprivation. This was supported by expression analyses of DR5uidA, root basipetal transport assay of auxin, and RSA of the pgp19 mutant exhibiting reduced auxin transport. A significant increase in the number of lateral roots under the Pi- Suc- condition in the chalcone synthase mutant (tt4-2) indicated a potential role for flavonoids in auxin-mediated Pi deficiency-induced modulation of RSA. The study thus demonstrated differential roles of Suc and auxin in the developmental responses of ontogenetically distinct root traits during Pi deprivation. In addition, lack of cross talk between local and systemic Pi sensing as revealed by the seedlings grown under either the Pi- Suc- condition or in the heterogeneous Pi environment highlighted the coexistence of Suc-independent and Suc-dependent regulatory mechanisms that constitute Pi starvation responses.     

Genetic analysis of adventitious root formation with a novel series of temperature-sensitive mutants of Arabidopsis thaliana

When cultured on media containing the plant growth regulator auxin, hypocotyl explants of Arabidopsis thaliana generate adventitious roots. As a first step to investigate the genetic basis of adventitious organogenesis in plants, we isolated nine temperature-sensitive mutants defective in various stages in the formation of adventitious roots: five root initiation defective (rid1 to rid5) mutants failed to initiate the formation of root primordia; in one root primordium defective (rpd1) mutant, the development of root primordia was arrested; three root growth defective (rgd1, rgd2, and rgd3) mutants were defective in root growth after the establishment of the root apical meristem. The temperature sensitivity of callus formation and lateral root formation revealed further distinctions between the isolated mutants. The rid1 mutant was specifically defective in the reinitiation of cell proliferation from hypocotyl explants, while the rid2 mutant was also defective in the reinitiation of cell proliferation from root explants. These two mutants also exhibited abnormalities in the formation of the root apical meristem when lateral roots were induced at the restrictive temperature. The rgd1 and rgd2 mutants were deficient in root and callus growth, whereas the rgd3 mutation specifically affected root growth. The rid5 mutant required higher auxin concentrations for rooting at the restrictive temperature, implying a deficiency in auxin signaling. The rid5 phenotype was found to result from a mutation in the MOR1/GEM1 gene encoding a microtubule-associated protein. These findings about the rid5 mutant suggest a possible function of the microtubule system in auxin response.     

Regulation of Shoot and Root Development through Mutual Signaling

Plants adjust their development in relation to the availability of nutrient sources. This necessitates signaling between root and shoot. Aside from the well-known systemic signaling processes mediated by auxin, cytokinin, and sugars, new pathways involving carotenoid-derived hormones have recently been identified. The auxin-responsive MAX pathway controls shoot branching through the biosynthesis of strigolactone in the roots. The BYPASS1 gene affects the production of an as-yet unknown carotenoid-derived substance in roots that promotes shoot development. Novel local and systemic mechanisms that control adaptive root development in response to nitrogen and phosphorus starvation were recently discovered. Notably, the ability of the NITRATE TRANSPORTER 1.1 to transport auxin drew for the first time a functional link between auxin, root development, and nitrate availability in soil. The study of plant response to phosphorus starvation allowed the identification of a systemic mobile miRNA. Deciphering and integrating these signaling pathways at the whole-plant level provide a new perspective for understanding how plants regulate their development in response to environmental cues.     

Extracellular ATP inhibits root gravitropism at concentrations that inhibit polar auxin transport

Raising the level of extracellular ATP to mM concentrations similar to those found inside cells can block gravitropism of Arabidopsis roots. When plants are grown in Murashige and Skoog medium supplied with 1 mM ATP, their roots grow horizontally instead of growing straight down. Medium with 2 mM ATP induces root curling, and 3 mM ATP stimulates lateral root growth. When plants are transferred to medium containing exogenous ATP, the gravity response is reduced or in some cases completely blocked by ATP. Equivalent concentrations of ADP or inorganic phosphate have slight but usually statistically insignificant effects, suggesting the specificity of ATP in these responses. The ATP effects may be attributable to the disturbance of auxin distribution in roots by exogenously applied ATP, because extracellular ATP can alter the pattern of auxin-induced gene expression in DR5-beta-glucuronidase transgenic plants and increase the response sensitivity of plant roots to exogenously added auxin. The presence of extracellular ATP also decreases basipetal auxin transport in a dose-dependent fashion in both maize (Zea mays) and Arabidopsis roots and increases the retention of [(3)H]indole-3-acetic acid in root tips of maize. Taken together, these results suggest that the inhibitory effects of extracellular ATP on auxin distribution may happen at the level of auxin export. The potential role of the trans-plasma membrane ATP gradient in auxin export and plant root gravitropism is discussed.     

The Control of Lateral Root Development in Cultured Pea Seedlings. II. Root Fasciation Induced by Auxin Inhibitors

Lateral root development in cultured seedlings of Pisum sativum (cv. Alaska) was modified by the application of auxin transport inhibitors or antagonists. When applied either to replace the root tip or beneath the cotyledonary node, two auxin transport inhibitors, 2,3,5-triiodobenzoic acid (TIBA) and 3,3a-dihydro-2-(p-methoxyphenyl)-8H-pyrazolo[5,1-α]isoindol-8-one (DPX-1840), increased cell division activity opposite the protoxylem poles. This resulted in the formation of masses of cells, which we are calling root primordial masses (RPMs), 2 to 3 days after treatment. RPMs differed from lateral root primordia in that they lacked apical organization. Some roots however developed both RPMs and lateral roots indicating that both structures were similar in terms of the timing and location of cell division in the pericycle and endodermis leading to their initiation. Removal of the auxin transport inhibitors allowed many of the RPMs to organize later into lateral root primordia and to emerge in clusters. When the auxin, indoleacetic acid (IAA) was added to the growth medium along with DPX-1840, 3 ranks of RPMs now in the form of fasciated lateral roots emerged from the primary root. The auxin antagonist, p-chlorophenoxy-isobutyric acid (PCIB), also induced RPM formation. In contrast to DPX-1840 treatment, the addition of IAA during PCIB treatment caused normal lateral root development.     

A role for auxin redistribution in the responses of the root system architecture to phosphate starvation in Arabidopsis

The changes in root system architecture (RSA) triggered by phosphate (P) deprivation were studied in Arabidopsis (Arabidopsis thaliana) plants grown for 14 d on 1 mM or 3 microM P. Two different temporal phases were observed in the response of RSA to low P. First, lateral root (LR) development was promoted between days 7 and 11 after germination, but, after day 11, all root growth parameters were negatively affected, leading to a general reduction of primary root (PR) and LR lengths and of LR density. Low P availability had contrasting effects on various stages of LR development, with a marked inhibition of primordia initiation but a strong stimulation of activation of the initiated primordia. The involvement of auxin signaling in these morphological changes was investigated in wild-type plants treated with indole-3-acetic acid or 2,3,5-triiodobenzoic acid and in axr4-1, aux1-7, and eir1-1 mutants. Most effects of low P on RSA were dramatically modified in the mutants or hormone-treated wild-type plants. This shows that auxin plays a major role in the P starvation-induced changes of root development. From these data, we hypothesize that several aspects of the RSA response to low P are triggered by local modifications of auxin concentration. A model is proposed that postulates that P starvation results in (1) an overaccumulation of auxin in the apex of the PR and in young LRs, (2) an overaccumulation of auxin or a change in sensitivity to auxin in the lateral primordia, and (3) a decrease in auxin concentration in the lateral primordia initiation zone of the PR and in old laterals. Measurements of local changes in auxin concentrations induced by low P, either by direct quantification or by biosensor expression pattern (DR5::beta-glucuronidase reporter gene), are in line with these hypotheses. Furthermore, the observation that low P availability mimicked the action of auxin in promoting LR development in the alf3 mutant confirmed that P starvation stimulates primordia emergence through increased accumulation of auxin or change in sensitivity to auxin in the primordia. Both the strong effect of 2,3,5-triiodobenzoic acid and the phenotype of the auxin-transport mutants (aux1, eir1) suggest that low P availability modifies local auxin concentrations within the root system through changes in auxin transport rather than auxin synthesis.     

Cytokinin Inhibits Lateral Root Development at the Earliest Stages of Lateral Root Primordium Initiation in Maize Primary Root

Root architecture is basically controlled by auxin and cytokinin, which antagonize in the formation of lateral roots (LRs) along the primary root (PR) axis. Several mechanisms have been proposed to explain the interaction between these two hormones, cytokinin being the hormone that inhibits LR formation. The analysis of the cytokinin effect on LR formation using LRs in several stages of development could indicate which steps of LR formation are more sensitive to cytokinin. The application of cytokinin to maize PRs showed that the inhibitory effect of cytokinin on LR formation was greater in the zones in which the initial events to form new LRs are taking place. In the presence of cytokinin, the PR is not able to produce new LRs in the initiation zone; this inhibitory effect is permanent as this zone did not recover the capability to form LRs after removing cytokinin. However, the LR density in zones with appreciable LR primordia when cytokinin was applied was only slightly inhibited when a high concentration was used. These results showed that LR formation is more sensitive to the inhibitory effect of cytokinin in the earliest stages of LR development. However, the elongation of a LR primordium to emerge and the subsequent elongation of the new LR were only slightly affected by cytokinin.

     

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.     

Light Quality Regulates Lateral Root Development in Tobacco Seedlings by Shifting Auxin Distributions

Light is an important environmental regulator of diverse growth and developmental processes in plants. However, the mechanisms by which light quality regulates root growth are poorly understood. We analyzed lateral root (LR) growth of tobacco seedlings in response to three kinds of light qualities (red, white, and blue). Primary (1°) LR number and secondary (2°) LR density were elevated under red light (on days 9 and 12 of treatment) in comparison with white and blue lights. Higher IAA concentrations measured in roots and lower in leaves of plants treated with red light suggest that red light accelerated auxin transport from the leaves to roots (in comparison with other light qualities). Corroborative evidence for this suggestion was provided by elevated DR5::GUS expression levels at the shoot/root junction and in the 2° LR region. Applications of N-1-naphthylphthalamic acid (NPA) to red light-treated seedlings reduced both 1° LR number and 2° LR density to levels similar to those measured under white light; DR5::GUS expression levels were also similar between these light qualities after NPA application. Results were similar following exogenous auxin (NAA) application to blue light-treated seedlings. Direct [³H]IAA transport measurement indicated that the polar auxin transport from shoot to root was increased by red light. Red light promoted PIN3 expression levels and blue light reduced PIN1, 3–4 expression levels in the shoot/root junction and in the root, indicating that these genes play key roles in auxin transport regulation by red and blue lights. Overall, our findings suggest that three kinds of light qualities regulate LR formation in tobacco seedlings through modification of auxin polar transport.     

Inhibition of Auxin Movement from the Shoot into the Root Inhibits Lateral Root Development in Arabidopsis

In roots two distinct polar movements of auxin have been reported that may control different developmental and growth events. To test the hypothesis that auxin derived from the shoot and transported toward the root controls lateral root development, the two polarities of auxin transport were uncoupled in Arabidopsis. Local application of the auxin-transport inhibitor naphthylphthalamic acid (NPA) at the root-shoot junction decreased the number and density of lateral roots and reduced the free indoleacetic acid (IAA) levels in the root and [³H]IAA transport into the root. Application of NPA to the basal half of or at several positions along the root only reduced lateral root density in regions that were in contact with NPA or in regions apical to the site of application. Lateral root development was restored by application of IAA apical to NPA application. Lateral root development in Arabidopsis roots was also inhibited by excision of the shoot or dark growth and this inhibition was reversible by IAA. Together, these results are consistent with auxin transport from the shoot into the root controlling lateral root development.     

Change in Size and Cell Number during the Development of Lateral Root Primordia in Zea mays L.

Increase in cell number, anlage volume and length have beeninvestigated during the overall period of lateral root primordiumdevelopment in excised primaries and in attached roots of Zeamays L. Each of these aspects of anlage growth was found toincrease more or less exponentially during the interval betweenprimordium initiation and subsequent emergence as a lateralin both batches of roots. Values were then determined for celldoubling time (Td), the size of the proliferative fraction (Pf),and for anlage volume (Tv) and length (Tt) doubling times duringthe overall period of primordium development and at intervalsduring this period in both the excised and attached roots. Thepattern of change which took place in Td, Tv, Tl and Pf duringlateral primordium development was found to be similar in bothbatches of roots. However, the overall period of anlage developmentwas shorter in the excised roots than in the attached ones.Moreover, when laterals grew out of the excised roots they didso with fewer cells than comparable laterals emerging from theattached roots. Zea mays L., maize, root primordia, lateral emergence, cell doubling time