Chemical root to shoot signaling under drought

Chemical signals are important for plant adaptation to water stress. As soils become dry, root-sourced signals are transported via the xylem to leaves and result in reduced water loss and decreased leaf growth. The presence of chemical signals in xylem sap is accepted, but the identity of these signals is controversial. Abscisic acid (ABA), pH, cytokinins, a precursor of ethylene, malate and other unidentified factors have all been implicated in root to shoot signaling under drought. This review describes current knowledge of, and advances in, research on chemical signals that are sent from roots under drought. The contribution of these different potential signals is discussed within the context of their role in stress signaling.     

干旱胁迫对甘薯苗期根系分化和生理特性的影响

以‘济薯26’和‘广薯87’为试验材料,采用砂培试验方法,以10%浓度的聚乙二醇(PEG-6000)模拟干旱胁迫,动态研究了干旱胁迫对甘薯发根分枝结薯期的3个不同阶段根系形态、内源激素、叶绿素荧光和活性氧代谢的影响.结果表明: 不同时期干旱胁迫均导致甘薯生物量显著降低,影响程度为移栽后第10天＞第20天＞第30天.与正常供水相比,移栽后10 d干旱胁迫导致甘薯平均直径和根体积下降幅度最大,其次是移栽后20 d,最后是移栽后30 d.双因素分析表明,不同时期和干旱胁迫显著影响了根系形态特征,且交互效应显著.不同时期干旱胁迫均显著降低了甘薯功能叶的叶绿素荧光特性,导致光合产物形成受阻,进而抑制了甘薯根系的分化.不同时期干旱胁迫均导致根系中促生长激素(吲哚乙酸和玉米素核苷)含量显著下降,抑制生长激素(脱落酸)含量显著上升,以及激素间比例失衡,从而抑制甘薯根系分化,且胁迫时间越早,甘薯根系分化受阻越严重.其中,济薯26根系分化受阻的严重程度显著低于广薯87.逐步回归分析表明,根内源激素和叶绿素荧光是影响平均直径和根体积的关键指标(R分别为0.936和0.972);通径分析表明,对甘薯平均直径和根体积的影响直接作用系数较大的是玉米素核苷、最大光化学效率和脱落酸.甘薯的水分临界期在发根分枝结薯期,其中移栽后第10天对干旱胁迫更敏感.在实际生产中,应加强栽苗后缓苗水分的供应.     

Identification of drought tolerance determinants by genetic analysis of root response to drought stress and abscisic Acid

Drought stress is a common adverse environmental condition that seriously affects crop productivity worldwide. Due to the complexity of drought as a stress signal, deciphering drought tolerance mechanisms has remained a major challenge to plant biologists. To develop new approaches to study plant drought tolerance, we searched for phenotypes conferred by drought stress and identified the inhibition of lateral root development by drought stress as an adaptive response to the stress. This drought response is partly mediated by the phytohormone abscisic acid. Genetic screens using Arabidopsis (Arabidopsis thaliana) were devised, and drought inhibition of lateral root growth (dig) mutants with altered responses to drought or abscisic acid in lateral root development were isolated. Characterization of these dig mutants revealed that they also exhibit altered drought stress tolerance, indicating that this root response to drought stress is intimately linked to drought adaptation of the entire plant and can be used as a trait to access the elusive drought tolerance machinery. Our study also revealed that multiple mechanisms coexist and together contribute to whole-plant drought tolerance.     

Field trials using single potato plants as whole plots, with and without nematicide treatment, to assess tolerance of attack by Globodera rostochiensis

It is important to assess the tolerance of new potato cultivars with partial resistance to potato cyst nematode attack. The experiments described here are intended to quantify tolerance adequately using a minimum of resources. When test plants were spaced widely in the field trials it was also possible to estimate their resistance levels but standard plant spacing allowed roots of neighbouring plants to encroach on a test plant's root space and to interfere with attempts to assay resistance. Standard plant spacing did, however, give more realistic yields per plant and competition from neighbouring plants was standardised by surrounding each test plant by four identical guard plants (cv. Désirée). Regressions of yield on severity of nematode attack were drawn when nematode infestation varied naturally across the trial site or when different levels of attack were created by applying nematicide at a range of dose rates. Cara was the most tolerant cultivar tested, with a complete spectrum of tolerance shown by other cultivars/clones. Some clones with a high degree of resistance were intolerant and it is precisely these that this type of test seeks to identify.     

小麦重组近交系群体株高和千粒重的抗旱遗传特性

以小麦重组近交系(RIL)群体(陇鉴19×Q9086)120个株系及其亲本为材料,研究雨养(DS)和灌溉(WW)条件下不同发育时期株高动态发育和千粒重的遗传特点及其相互关系,并且评价群体性状的遗传变异.结果表明:在2种水分条件下,小麦RIL群体表型变异广泛且存在超亲分离,对水分环境敏感,拔节期株高具有较高的旱胁迫系数(DSC=0.851).不同发育时期株高与千粒重呈显著正相关,与其他时期相比,拔节期株高与千粒重有较高的相关系数(R2DS=0.32,R2ww=0.28).拔节期株高和开花期株高对千粒重有显著的正向直接作用,但间接总效应为负向作用;抽穗期株高和成熟期株高的作用与此相反.各性状遗传力普遍较低,为0.27 ～0.60;在雨养和灌溉条件下,控制千粒重的基因对数分别为10和13,控制株高的基因对数分别为3～7和4～ 14.表型DSC聚类将小麦RIL群体分为5类,群体内表型和对水分敏感程度变异丰富,该群体适合进行抗旱性状数量遗传研究.     

Physiological and molecular analysis on root growth associated with the tolerance to aluminum and drought individual and combined in Tibetan wild and cultivated barley

Planta - The drought-stimulated gene expression of NCED, SUS, and KS - DHN and ABA signal cross-talk with other phytohormones maintains barley root growth under drought stress at pH 4.0 plus...     

Low leaf hydraulic conductance associated with drought tolerance in soybean

Lack of water is the most serious environmental constraint on agricultural production. More efficient use of water resources is a key solution for increased plant productivity in water-deficit environments. We examined the hydraulic characteristics of a 'slow wilting' phenotype in soybean ( Glycine max Merr.), PI 416937, which has been shown to have relatively constant transpiration rates above a threshold atmospheric vapor pressure deficit (VPD). The VPD response of PI 416937 was confirmed. Three experiments are reported to examine the hypothesis that the VPD response was a result of low hydraulic conductance in leaves as compared to two other soybean genotypes. Results are reported from experiments to measure transpiration response to VPD when xylem water potential was maintained at zero, leaf rehydration response and leaf carbon assimilation response to petiole cutting. Major interspecific differences in leaf hydraulic properties were observed. The observed low leaf hydraulic conductance in PI 416937 is consistent with an increased water use efficiency, and an increased water conservation by limiting transpiration rates under high evaporative conditions but allowing normal gas exchange rates under more moderate evaporative conditions.     

Whole-plant carbon relations and root respiration associated with root tolerance to high soil temperature for Agrostis grasses

Plant tolerance to high soil temperature may be related to the adjustment in carbon production and utilization. The objective of this study was to determine changes in whole-plant carbon balance and root respiration rate in relation to root tolerance to high soil temperature for two Agrostis grass species varying in heat tolerance. Plant tolerance to high soil temperature was compared between Agrostis scabra, a thermal grass species adapted to chronic high-temperature soils in the geothermal areas in Yellowstone National Park, and creeping bentgrass (Agrostis stolonifera), a cultivated grass species adapted to cool climatic regions. Plant roots were exposed to low soil temperature (20 °C) or high soil temperature (37 °C) for 17 days in water baths placed in a controlled-environment growth chamber. Root biomass and cell membrane stability were determined to evaluate root thermotolerance of both species. Canopy photosynthetic rate (Pn), whole-plant respiration rate, root respiration rate, and total non-structural carbohydrate (TNC) content were measured to assess changes in carbon production and utilization in response to high soil temperature. Root biomass and cell membrane stability declined with increasing soil temperature, but the decline was much less for A. scabra than A. stolonifera, suggesting that roots of A. scabra were more tolerant to heat stress. Canopy Pn decreased and whole-plant respiration rate increased for A. stolonifera, but canopy Pn and respiration rate were unchanged for A. scabra in response to increasing soil temperature. After 17 days of high soil temperature treatment, A. stolonifera exhibited carbon deficit at the whole-plant level, whereas A. scabra maintained positive carbon gain. Root respiration of plants previously grown at 20 °C increased after a short-term treatment (24 h) at 37 °C, but the increase was significantly lower for A. scabra than for A. stolonifera. TNC content in roots did not show response to short-term (24 h) changes in temperature and did not exhibit species variations. Leaves of A. scabra, however, maintained TNC content under both low and high temperature regimes. Our results suggest that root thermotolerance of cool-season grasses could be related to the maintenance of positive whole-plant carbon balance, and down-regulation of whole-plant and root respiration rates in response to increasing soil temperature.     

Enhanced drought and salinity tolerance in transgenic potato plants with a BADH gene from spinach

Drought and salinity are the most important abiotic stresses that affect the normal growth and development of plants. Glycine betaine is one of the most important osmolytes present in higher plants that enable them to cope with environmental stresses through osmotic adjustment. In this study, a betaine aldehyde dehydrogenase (BADH) gene from spinach under the control of the stress-induced promoter rd29A from Arabidopsis thaliana was introduced into potato cultivar Gannongshu 2 by the Agrobacterium tumefaciens system. Putative transgenic plants were confirmed by Southern blot analysis. Northern hybridization analysis demonstrated that expression of BADH gene was induced by drought and NaCl stress in the transgenic potato plants. The BADH activity in the transgenic potato plants was between 10.8 and 11.7 U. There was a negative relationship (y = −2.2083x + 43.329, r = 0.9495) between BADH activity and the relative electrical conductivity of the transgenic potato plant leaves. Plant height increased by 0.4–0.9 cm and fresh weight per plant increased by 17–29% for the transgenic potato plants under NaCl and polyethylene glycol stresses compared with the control potato plants. These results indicated that the ability of transgenic plants to tolerate drought and salt was increased when their BADH activity was increased.     

不同抗旱性花生品种的根系形态发育及其对干旱胁迫的响应

为明确不同抗旱性花生品种的根系形态发育特征,探讨其根系形态发育特征对不同土壤水分状况的响应机制,在防雨棚旱池内进行土柱栽培试验,研究抗旱型品种“花育22号”、“唐科8号”和干旱敏感型品种“花育23号”3个不同抗旱性花生品种根系形态发育特征及其对干旱胁迫的响应.结果表明:抗旱型品种根系较发达,具有较大的根系生物量、总根长、总根系表面积.干旱胁迫使抗旱型品种根系总表面积和体积增加,而干旱敏感型品种则相反.干旱胁迫显著增加抗旱型品种“花育22号”20 cm以下土层内根长密度分布比例及根系表面积和体积,但“唐科8号”相应根系性状仅在20-40 cm土层内增加;干旱胁迫使干旱敏感型品种“花育23号”40 cm以下土层内各根系性状升高,但未达显著水平且其深层土壤内各根系性状增加幅度小于“花育22号”.花生根系总长、总表面积及0-20 cm土层内根系性状与产量间呈显著或极显著正相关.土壤水分亏缺条件下,花生主要通过增加深层土壤内根长、根系表面积和体积等形态特性,优化空间分布构型,以调节植株对水分的利用.     

Field performance of transgenic potato plants compared with controls regenerated from tuber discs and shoot cuttings

Summary The objective of this study was to separate and determine effects on the field performance of transgenic potatoes that originate from the tissue culture process of transformation and from the genes inserted. The constructs introduced contained the reporter gene for betaglucuronidase (GUS) under the control of the patatin promoter (four different constructs) and the neomycin phosphotransferase gene under the control of the nopaline synthase promoter. Both genes might be expected to have a neutral effect on plant phenotype. The field performance of transgenic plants (70 independent transformants) was compared with non-transgenic plants regenerated from tuber discs by adventitious shoot formation and from shoot cultures established from tuber nodal cuttings. Plants from all three treatments were grown in a field trial from previously field-grown tubers, and plant performance was measured in terms of plant height at flowering, weight of tubers, number of tubers, weight of large tubers and number of large tubers. There was evidence of somaclonal variation among the transgenic plants; mean values for all characters were significantly lower and variances generally higher than from plants derived from nodal shoot cultures. A similar change in means and variances was observed for the non-transgenic tuber-disc regenerants when compared with shoot culture plants. Plant height, tuber weight and tuber number were, however, significantly lower in transgenic plants than in tuber-disc regenerants, suggesting an effect on plant performance either of the tissue culture process used for transformation or of the genes inserted. There were significant differences between constructs for all five plant characters. The construct with the smallest segment of patatin promoter and the lowest level of tuber specificity for GUS expression had the lowest values for all five characters. It is proposed that the nature of GUS expression is influencing plant performance. There was no indication that the NPTII gene, used widely in plant transformation, has any substantial effect on plant performance in the field.     

Protein storage and root:shoot reallocation provide tolerance to damage in a hybrid willow system

To determine the mechanistic basis of tolerance, we evaluated six candidate traits for tolerance to damage using F₂ interspecific hybrids in a willow hybrid system. A distinction was made between reproductive tolerance and biomass tolerance; reproductive tolerance was designated as a plant’s proportional change in catkin production following damage, while biomass tolerance referred to a plant’s proportional change in biomass (i.e., regrowth) following damage. F₂ hybrids were generated to increase variation and independence among candidate traits. Using three clonally identical individuals, pre-damage candidate traits for tolerance to damage (root:shoot ratio, total nonstructural carbohydrate, and total available protein) and post-damage candidate traits (relative root:shoot ratio, phenolic ratio, and specific leaf area ratio) were measured. The range of variation for these six candidate traits was broad. Biomass was significantly increased two years after 50% shoot length removal, and catkin production was not significantly reduced when damaged, suggesting that F₂ hybrids had great biomass tolerance and reproductive tolerance. Based on multiple regression methods, increased reproductive tolerance was associated with increased protein storage and decreased relative root:shoot ratio (reduced root allocation after damage). In addition, a positive relationship between biomass tolerance and condensed tannins was detected, and both traits were associated with increased reproductive tolerance. These four factors explained 57% of the variance in the reproductive tolerance of F₂ hybrids, but biomass tolerance explained the majority of the variance in reproductive tolerance. Changes in plant architecture in response to plant damage may be the underlying mechanism that explains biomass tolerance.     

干旱胁迫对不同耐旱性玉米杂交种产量和根系生理特性的影响

通过盆栽人工模拟干旱试验,研究了全生育期中度干旱胁迫对不同耐旱性玉米杂交种（耐旱：京科628;不耐旱：农大95）产量及根系生理特性的影响.结果表明：干旱胁迫下,耐旱玉米产量比对照减少33.7%,不耐旱玉米则比对照减少62.3%.干旱胁迫下,玉米根系生物量降低且最大值出现时间提前,与对照相比,不耐旱玉米根冠比升高,耐旱玉米根冠比前期升高后期降低;根系活力降低,不耐旱玉米根系活力降低幅度大于耐旱玉米;根系超氧化物歧化酶（SOD）活性前期高于对照后期低于对照,耐旱玉米根系SOD活性开始低于对照的时间比不耐旱玉米晚;根系丙二醛（MDA）含量升高,随干旱胁迫处理时间的延长,不耐旱玉米MDA含量比对照升高的幅度大于耐旱玉米;根系可溶性蛋白含量降低,不耐旱玉米的降低幅度大于耐旱玉米.干旱胁迫下耐旱玉米杂交种根系活力﹑根系SOD活性及可溶性蛋白含量较高,减缓了根系的衰老进程,延长了根系功能期,这可能是耐旱玉米杂交种在干旱胁迫下仍能获得较高产量的重要原因之一.     

两个不同耐旱性棉花品种根系生理特性对干旱的响应

为探明不同抗旱性棉花品种的根系生理特性对干旱的响应及其与生物量的关系,以不耐旱性品种‘新陆早17号'(L17)和耐旱性品种‘新陆早22号'(L22)为试材,在土柱栽培条件下设常规灌溉(CK)、轻度干旱(W₁)和中度干旱(W₂)处理,研究干旱胁迫对不同耐旱性棉花品种根系活力、保护酶活性及解剖结构(导管直径、数量)和生物量的影响。结果表明:干旱胁迫下两棉花品种根系可溶性蛋白(SP)含量、根系活力(RV)、木栓层数、根茎导管数量、导管直径显著降低,根系丙二醛(MDA)含量及保护性酶活性显著增加,进而导致地上部干物质量显著降低。与L17相比,L22的SP含量、0～40 cm和80～120 cm土层RV、木栓层数、根茎导管数量、导管直径以及地上部干物质量均显著增加,尤其W₂条件下L22的RV降幅比L17低26.2%,过氧化氢酶(CAT)、过氧化物酶(POD)和超氧化物歧化酶(SOD)活性和皮层厚度分别比L17高43.6%、6.9%、25.4%、19.9%,且差异均达到显著水平。干物质量与RV、SOD、POD、木栓层数、导管直径和个数之间均呈显著正相关。因此,耐旱性强的棉花品种在干旱条件下通过保持较高的根系活性、木栓层数、导管直径和数量,进而促进地上部生物量积累,是其具有较高耐旱性的生理机制。     

Genetic variation in ZmTIP1 contributes to root hair elongation and drought tolerance in maize

Drought is a major abiotic stress that threatens maize production globally. A previous genome‐wide association study identified a significant association between the natural variation of ZmTIP1 and the drought tolerance of maize seedlings. Here, we report on comprehensive genetic and functional analysis, indicating that ZmTIP1, which encodes a functional S‐acyltransferase, plays a positive role in regulating the length of root hairs and the level of drought tolerance in maize. We show that enhancing ZmTIP1 expression in transgenic Arabidopsis and maize increased root hair length, as well as plant tolerance to water deficit. In contrast, ZmTIP1 transposon‐insertional mutants displayed the opposite phenotype. A calcium‐dependent protein kinase, ZmCPK9, was identified as a substrate protein of ZmTIP1, and ZmTIP1‐mediated palmitoylation of two cysteine residues facilitated the ZmCPK9 PM association. The results of this research enrich our knowledge about ZmTIP1‐mediated protein S‐acylation modifications in relation to the regulation of root hair elongation and drought tolerance. Additionally, the identification of a favourable allele of ZmTIP1 also provides a valuable genetic resource or selection target for the genetic improvement of maize.     

Locating genes associated with root morphology and drought avoidance in rice via linkage to molecular markers

This research was undertaken to identify and map quantitative trait loci (QTLs) associated with five parameters of rice root morphology and to determine if these QTLs are located in the same chromosomal regions as QTLs associated with drought avoidance/tolerance. Root thickness, root:shoot ratio, root dry weight per tiller, deep root dry weight per tiller, and maximum root length were measured in three replicated experiments (runs) of 203 recombinant inbred lines grown in a greenhouse. The lines were from a cross between indica cultivar Co39 andjaponica cultivar Moroberekan. The 203 RI lines were also grown in three replicated field experiments where they were drought-stressed at the seedling, early vegetative, and late-vegetative growth stage and assigned a visual rating based on leaf rolling as to their degree of drought avoidance/tolerance. The QTL analysis of greenhouse and field data was done using single-marker analysis (ANOVA) and interval analysis (Mapmaker QTL). Most QTLs that were identified were associated with root thickness, root/shoot ratio, and root dry weight per tiller, and only a few with deep root weight. None were reliably associated with maximum root depth due to genotype-by-experiment interaction. Root thickness and root dry weight per tiller were the characters found to be the least influenced by environmental differences between greenhouse runs. Correlations of root parameters measured in greenhouse experiments with field drought avoidance/tolerance were significant but not highly predictive. Twelve of the fourteen chromosomal regions containing putative QTLs associated with field drought avoidance/tolerance also contained QTLs associated with root morphology. Thus, selecting for Moroberekan alleles at marker loci associated with the putative root QTLs identified in this study may be an effective strategy for altering the root phenotype of rice towards that commonly associated with drought-resistant cultivars.     

Accumulation of terpenoid phytoalexins in maize roots is associated with drought tolerance

Maize (Zea mays) production, which is of global agro‐economic importance, is largely limited by herbivore pests, pathogens and environmental conditions, such as drought. Zealexins and kauralexins belong to two recently identified families of acidic terpenoid phytoalexins in maize that mediate defence against both pathogen and insect attacks in aboveground tissues. However, little is known about their function in belowground organs and their potential to counter abiotic stress. In this study, we show that zealexins and kauralexins accumulate in roots in response to both biotic and abiotic stress including, Diabrotica balteata herbivory, Fusarium verticillioides infection, drought and high salinity. We find that the quantity of drought‐induced phytoalexins is positively correlated with the root‐to‐shoot ratio of different maize varieties, and further demonstrate that mutant an2 plants deficient in kauralexin production are more sensitive to drought. The induction of phytoalexins in response to drought is root specific and does not influence phytoalexin levels aboveground; however, the accumulation of phytoalexins in one tissue may influence the induction capacity of other tissues.