小麦根系特征对干旱胁迫的响应

干旱胁迫时, 小麦(Triticum aestivum)根系率先产生应激响应, 同时向地上部发出信号, 诱导地上部发生生理反应, 从而提高植株抗旱能力。根系构型包括平面几何性状和立体几何结构(即拓扑构型), 具有遗传稳定性和可塑性。干旱胁迫影响根系理化特性, 如根源化学信号、根系细胞酶类和根系渗透作用的响应。根系通过调整其解剖学结构和水分吸收动力等来适应干旱胁迫。该文从根系构型、理化特性和解剖学结构3个方面, 系统阐述了小麦根系特征对干旱胁迫的响应, 并探讨了其与干旱胁迫的关系和当前研究中存在的问题, 以期为相关研究提供参考。     

小麦根系与土壤水分胁迫关系的研究进展

几十年来,大量科学工作者为拓宽小麦根系对土壤水分的吸收能力和调控根系对干旱的适应能力,挖掘干旱地区的生产潜力,实现高产做了大量细致的研究工作,取得了许多重要研究成果,综述了土壤水分胁迫对小麦根系形态、构型建成和生理指标影响的影响。过去进行的研究表明,干旱胁迫条件下,不仅表达小麦根系形态和构型建成指标的根系数量、根系比表面积、根冠比、根生长势、根水势,导管直径等发生显著变化,而且表达根系生理指标的伤流流、根呼吸速率、根系质膜透性、膜脂过氧化水平、保护酶及其同工酶等也发生相应改变,虽然不同的研究者所获得的研究结果不同,有的甚至相互矛盾,但从总体看,各种变要是对干旱胁迫的一种适应性反应,有利于提高小麦的抗旱能力,对干旱条件下产量的形成具有重要作用。     

盐旱复合胁迫对小麦幼苗生长和水分吸收的影响

为明确盐害、干旱及盐旱复合胁迫对小麦幼苗生长和水分吸收的影响,从而为盐害和干旱胁迫下栽培调控提供理论依据。以2个抗旱性不同的小麦品种(扬麦16和耐旱型洛旱7号)为材料,采用水培试验,以NaCl和PEG模拟盐旱复合胁迫,研究了盐旱复合胁迫下小麦幼苗生长、根系形态、光合特性及水分吸收特性的变化。结果表明,盐、旱及复合胁迫下小麦幼苗的生物量、叶面积、总根长与根系表面积、叶绿素荧光和净光合速率均显著下降,但是复合胁迫处理的降幅却显著低于单一胁迫。盐旱复合胁迫下根系水导速率和根系伤流液强度显著大于单一胁迫,从而提高了小麦幼苗叶片水势和相对含水量。盐胁迫下小麦幼苗Na~+/K~+显著大于复合胁迫,但复合胁迫下ABA含量却显著小于单一的盐害和干旱胁迫。因此,盐旱复合胁迫可以通过增强根系水分吸收及降低根叶中ABA含量以维持较高光合能力,这是盐旱复合胁迫提高小麦适应性的重要原因。洛旱7号和扬麦16对盐及盐旱复合胁迫的响应基本一致,但在干旱胁迫下洛旱7号表现出明显的耐性。     

干旱胁迫对小麦幼苗根系生长和叶片光合作用的影响

采用水培试验方法,以2个耐旱性不同的小麦品种(敏感型望水白和耐旱型洛旱7号)为材料,研究了干旱胁迫对小麦幼苗根系形态、生理特性以及叶片光合作用的影响,以期揭示小麦幼苗对干旱胁迫的适应机制.结果表明： 干旱胁迫下,2个小麦品种幼苗的根系活力显著增大,而根数和根系表面积受到抑制;干旱胁迫降低了望水白的叶片相对含水量,提高了束缚水/自由水,而对洛旱7号无显著影响;干旱胁迫降低了2个小麦品种叶片的叶绿素含量、净光合速率、蒸腾速率、气孔导度和胞间CO2浓度,但随胁迫时间的延长,洛旱7号的叶绿素含量和净光合速率与对照差异不显著;干旱胁迫降低了2个小麦品种幼苗的单株叶面积,以及望水白的根系、地上部和植株生物量,而对洛旱7号无显著影响.水分胁迫下,耐旱型品种可以通过提高根系活力、保持较高的根系生长量来补偿根系吸收面积的下降,保持较高的根系吸水能力,进而维持较高的光合面积和光合速率,缓解干旱对生长的抑制.     

干旱胁迫对小麦幼苗根系生长和叶片光合作用的影响

采用水培试验方法,以2个耐旱性不同的小麦品种(敏感型望水白和耐旱型洛旱7号)为材料,研究了干旱胁迫对小麦幼苗根系形态、生理特性以及叶片光合作用的影响,以期揭示小麦幼苗对干旱胁迫的适应机制.结果表明： 干旱胁迫下,2个小麦品种幼苗的根系活力显著增大,而根数和根系表面积受到抑制;干旱胁迫降低了望水白的叶片相对含水量,提高了束缚水/自由水,而对洛旱7号无显著影响;干旱胁迫降低了2个小麦品种叶片的叶绿素含量、净光合速率、蒸腾速率、气孔导度和胞间CO2浓度,但随胁迫时间的延长,洛旱7号的叶绿素含量和净光合速率与对照差异不显著;干旱胁迫降低了2个小麦品种幼苗的单株叶面积,以及望水白的根系、地上部和植株生物量,而对洛旱7号无显著影响.水分胁迫下,耐旱型品种可以通过提高根系活力、保持较高的根系生长量来补偿根系吸收面积的下降,保持较高的根系吸水能力,进而维持较高的光合面积和光合速率,缓解干旱对生长的抑制.     

胡杨幼苗根系生长与构型对土壤水分的响应

胡杨实生幼苗成活率低是制约其更新与人工育苗保存的关键问题,而幼苗根系生长与构型是影响其存活的重要因素。该试验以1年生胡杨幼苗为材料,通过2种给水方式(断续给水和连续给水)下各6个土壤水分梯度处理的控制试验,探究胡杨幼苗根系生长与构型对荒漠地区关键因子水分的响应特征。结果表明:(1)2种给水方式下干旱胁迫均使根冠比增加,且断续给水处理下幼苗根冠比显著大于连续给水。(2)一定程度的干旱处理还可以促进幼苗根系形态发育特征的发展和根系生物量的积累,但过度干旱胁迫或土壤水分含量过多都会抑制根系生长,并以连续给水、土壤含水量15%处理下幼苗根系最为发达。(3)幼苗深扎根能力强,其根宽深比在2种给水方式下均小于1,且断续给水处理显著小于连续给水处理;2种给水方式下根宽深比都与土壤水分含量呈显著正相关。(4)根系拓扑指数在2种给水方式下无显著差异且均接近1,但都与土壤水分含量呈显著负相关。即幼苗根系趋向鱼尾状分支结构,次级分支少,这种根系延伸策略有利于胡杨幼苗在干旱贫瘠的土壤环境中生存。(5)根系构型参数的主成分分析显示,总根长、总根表面积、根宽深比和拓扑指数在2种给水方式下都能很好地表示胡杨幼苗根系构型特征。可见,胡杨幼苗根系通过构筑鱼尾状分支结构、增加垂直根纵向延伸能力和增大根冠比适应干旱环境;水分对于胡杨幼苗根系生长与构型作用显著,根系对水分因子的响应对于胡杨幼苗适应水分异质性环境具有重要意义。     

干旱胁迫对不同烤烟品种根系生长和生理特性的影响

以抗旱型烤烟品种‘农大202'及一般型烤烟品种‘NC89'和‘K326'为材料,采用盆栽试验研究了干旱胁迫对根系生长和生理特性的影响,以明确各烟草品种的抗旱能力及其与根系生长和生理特性的关系.结果显示:严重干旱胁迫之后各烤烟品种根系鲜重、干重上升,而根系活力、根系吸收面积以及根系SOD和POD活性等根系生理指标则呈下降趋势.在干旱胁迫条件下,‘农大202'的根系总吸收面积、活跃吸收面积、根系活力、SOD活性和POD活性均显著高于两对照品种‘NC89'和‘K326',但其根系鲜重和干重等生物量并不具有太大的优势.研究表明,烟草的根系生长和生理特性对环境水分条件的响应存在明显的基因型差异;在干旱胁迫条件下, ‘农大202'根系能保持较高总吸收面积、活跃吸收面积、根系活力和保护酶活性,是其具有较强抗旱性的生理基础.     

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.     

干旱胁迫对花生根系生长发育和生理特性的影响

以花育17号和唐科8号两个花生品种为试验材料,在防雨棚栽培池内进行土柱栽培试验,研究了中度干旱胁迫和正常供水处理下花生生育后期根系形态发育特征和生理特性.结果表明: 唐科8号具有较发达的根系及较高的产量和抗旱系数,花育17号根系对干旱胁迫的适应性小于唐科8号.两品种根长密度、根系生物量均主要分布于0～40 cm土层中,但同一土层内两品种根系性状存在差异.与正常供水处理相比,干旱胁迫处理使花育17号各生育期总根长、根系总表面积和总体积均降低,而唐科8号除花针期显著降低外,其余生育期均明显升高;干旱胁迫增加了两品种20～40 cm土层内根系生物量、根系表面积和体积,而降低了40 cm以下土层内各根系性状;干旱胁迫处理使两品种饱果期40 cm以下土层内根系活力降低,且花育17号降低幅度高于唐科8号.干旱胁迫下两品种生育后期根系发育和生理特性的差异表明其根系在干旱胁迫下对水分吸收和利用存在差异.     

小麦ARF基因家族生物信息学分析及在干旱胁迫下的表达特性研究

植物生长素响应因子ARF(auxin response factor)参与调节了植物的向性运动、顶端优势、微观的分化、侧根和茎的形态发生等众多生理反应,在植物生长发育的整个过程都起到重要调控作用。本研究通过对小麦最新基因组数据进行分析,获得了61个ARF家族基因,命名为TaARFs,根据染色体编号排列为TaARF1~TaARF61,对61个TaARFs基因进行系统生物信息学分析后发现ARF家族基因结构较为复杂,外显子数量从1个到15个变化不等,除了4号染色体和5A和5B染色体之外,其余的染色体均有ARF家族基因分布。ARF家族基因大多包含B3 DNA结构域、ARF结构域(Auxin-resp)和Aux/IAA结构域;同源进化分析表明,小麦ARF家族基因的旁系同源基因数量明显多于大麦和二穗短柄草。通过拟南芥数据库比对获得14个高同源的根系发育相关的小麦ARF家族基因,利用二系杂交小麦京麦6号及父母本根系为试材进行干旱胁迫处理及实时荧光定量PCR(qPCR)筛选。结果表明,7个小麦ARF基因不同程度受到干旱胁迫诱导,其在旱胁迫下的表达量显著高于正常条件下的表达量,可能参与干旱胁迫应答;此外本研究还发现,ARF基因在F1杂交种中表达量显著高于双亲,表现出超亲表达模式,可能参与了根系抗旱杂种优势基因表达调控网络。     

Quantifying relationships between rooting traits and water uptake under drought in Mediterranean barley and durum wheat

In Mediterranean regions drought is the major factor limiting spring barley and durum wheat grain yields. This study aimed to compare spring barley and durum wheat root and shoot responses to drought and quantify relationships between root traits and water uptake under terminal drought.One spring barley（Hordeum vulgare L. cv. Rum） and two durum wheat Mediterranean cultivars（Triticum turgidum L. var durum cvs Hourani and Karim） were examined in soil‐column experiments under well watered and drought conditions. Root system architecture traits, water uptake, and plant growth were measured. Barley aerial biomass and grain yields were higher than for durum wheat cultivars in well watered conditions. Drought decreased grain yield more for barley（47%） than durum wheat（30%, Hourani）. Root‐to‐shoot dry matter ratio increased for durum wheat under drought but not for barley, and root weight increased for wheat in response todrought but decreased for barley. The critical root length density（RLD） and root volume density（RVD） for 90% available water capture for wheat were similar to（cv. Hourani） or lower than（cv. Karim） for barley depending on wheat cultivar. For both species, RVD accounted for a slightly higher proportion of phenotypic variation in water uptake under drought than RLD.     

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.     

植物功能性状对土壤保持的影响研究述评

植被对土壤保持具有重要的影响,但是从植物功能性状的角度总结评述植被对土壤保持影响的研究并不多见。总结评述了植物地上功能性状、地下功能性状对土壤保持功能的影响以及植物地上、地下功能性状的关系,认为:(1)植被地上部分功能性状对土壤保持的作用主要体现在对溅蚀、面蚀的影响及间接改变土壤理化性质等方面,其功能性状指标主要包括叶面积、叶长、叶宽、枝数、植被高度等;(2)植被地下部分功能性状对土壤保持的作用主要体现在固持土壤、提高土壤抗剪切强度、提高土壤抗侵蚀能力、增强土壤渗透性,植物根系固持土壤与根系抗拉能力密切相关,植物根系土壤的物理和水文性质,与细根比例、根长密度、根表面积等性状密切相关;(3)可以通过植物地上部分功能性状间接反映地下部分功能性状,但是现有研究多为定性认识;(4)在植物功能性状对土壤保持的研究中亟待加强植被地上地下功能性状的长期定位监测,深化植被功能性状尤其是根系特征与土壤保持的作用机理,加强植被地上部分、地下部分功能性状的定量表达,建立植被功能性状与土壤保持功能的定量关系,实现植被功能性状与土壤保持功能特征的动态链接。     

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。     

Correlation of drought traits and the predictability of osmotic potential at full leaf turgor in vegetation from New Zealand

Scientists do not know precisely how severe will be the impact of climate change on species. Evidence suggests that for some species, their future distributions might be jeopardized by local extinctions and drought‐induced tree mortality. Thus, we require models capable of estimating drought tolerance across many species. We can approach this goal by assessing functional traits. The trait osmotic potential at full turgor, π_O, is potentially a good drought indicator; however, few studies address its importance as a drought‐tolerance predictor and it is difficult to measure in the field with accuracy. In this work, we aim to answer the questions: which drought traits correlate with π_O?; do morpho‐anatomical traits correlate with π_O?; and which trees and shrubs are more (or less) vulnerable to drought? To achieve this aim, we assessed physiological and morpho‐anatomical traits for 14 native species from New Zealand forests. We included leaf‐ and wood‐related traits, π_O, water potential and stomatal conductance. We examined how these traits correlate with π_O and sought to generate models to predict π_O as a function of other traits. We tested 33 different models and evaluated them using Akaike's information criterion. Unfortunately, none of the morpho‐anatomical traits correlated well with π_O. Instead, water potential correlated most strongly with π_O. None of the models using only morpho‐anatomical traits produced plausible results. The model with the best predictive performance incorporated the effects of both morpho‐anatomical and physiological traits: water potential and wood saturated water content. Of the species analysed, and based on their π_O response, Lophozonia menziesii was considered the most vulnerable to drought stress, whereas Plagianthus regius was the least vulnerable. Our findings imply that it is potentially valuable to keep exploring the use of π_O as a drought indicator and that the effort required to measure some physiological traits, such as water potential, may be essential to consider plant drought responses and to predict π_O.     

A meta-analysis on morphological,physiological and biochemical responses of plants with PGPR inoculation under drought stress

Plant growth-promoting rhizobacteria (PGPR) can help plants to resist drought stress. However, the mechanisms of how PGPR inoculation affect plant status under drought remain incompletely understood. We performed a meta-analysis of plant response to PGPR inoculation by compiling data from 57 PGPR-inoculation studies, including 2, 387 paired observations on morphological, physiological and biochemical parameters under drought and well-watered conditions. We compare the PGPR effect on plants performances among different groups of controls and treatments. Our results reveal that PGPR enables plants to restore themselves from drought-stressed to near a well-watered state, and that C4 plants recover better from drought stress than C3 plants. Furthermore, PGPR is more effective underdrought than well-watered conditions in increasing plant biomass, enhancing photosynthesis and inhibiting oxidant damage, and the responses of C4 plants to the PGPR effect was stronger than that of C3 plants under drought conditions. Additionally, PGPR belonging to different taxa and PGPR with different functional traits have varying degrees of drought-resistance effects on plants. These results are important to improve our understanding of the PGPR beneficial effects on enhanced drought-resistance of plants.     

模拟氮沉降和干旱对准噶尔盆地两种一年生荒漠植物生长和光合生理的影响

氮素和水分是荒漠生态系统的两个主要限制因子, 研究两者对荒漠植物的效应有助于深入了解荒漠生态系统对全球变化的响应。该文选择准噶尔盆地荒漠地区两种常见的一年生植物涩荠(Malcolmia africana)和钩刺雾冰藜(Bassia hyssopifolia), 设置0、0.18和0.72 g N·m ^-2·week ^-13个施氮浓度和湿润与干旱两个土壤水分处理, 研究模拟氮沉降增加和干旱对其生长和光合生理的影响。结果表明: (1)两种植物的根长、根重、叶片数、叶面积、总生物量和冠根比均随着施氮浓度的增加而增加, 干旱能够抑制氮对植物生长的促进作用, 但是, 氮的增加同时也能部分缓解干旱对植物生长的影响。与钩刺雾冰藜相比, 涩荠的根长、生物量和冠根比更易受氮增加和干旱的影响。(2)两种植物的最大净光合速率、叶绿素含量、可溶性蛋白含量随着氮浓度增加而增加, 但涩荠和钩刺雾冰藜对氮增加和干旱的生理响应也有所不同, 涩荠的响应更加敏感。两种植物对氮沉降和干旱胁迫响应的差异可能是其生活型等生物学特性差异所引起。通过对两种一年生植物的生长和光合生理分析表明, 在古尔班通古特沙漠, 春季丰富的降水和氮素增加将有利于涩荠和钩刺雾冰藜的生长和生产力的增加, 相对地下生长, 地上部分增加更显著。当干旱季节来临时, 氮的增加又能够在一定程度上降低干旱对这两种植物的负效应, 说明其对干旱具有一定的生态补偿作用。