不同磷水平下小麦-蚕豆间作根系形态的变化及其与内源激素的关系

采用水培方法,研究了不同磷水平下小麦-蚕豆间作体系根系形态变化及其与内源激素的相关关系。结果表明: 与单作小麦相比,在低磷(1/2P)水平下,小麦-蚕豆间作能显著增加小麦的根长,显著减少小麦根系的平均直径,显著增加根系的表面积;在常规磷(P)水平下,间作能显著降低小麦根系的平均直径,有增加小麦根长和根表面积的趋势;与单作蚕豆相比,间作能明显促进蚕豆根系的增长,同时增加蚕豆根表面积。在1/2P水平下,间作能显著提高小麦和蚕豆根系中的生长素(IAA)、脱落酸(ABA)、水杨酸(SA)和茉莉酸(JA)含量;在P水平下,间作能显著提高小麦根系中的IAA、ABA和JA含量,单、间作小麦根系中的SA含量没有显著差异,间作显著增加了蚕豆根系中ABA和SA含量,单、间作蚕豆根系中的IAA和JA含量无显著差异。单作条件下,小麦和蚕豆根系中的内源激素(IAA、ABA、SA和JA)含量与其根系形态(根长、根平均直径和根表面积)无显著相关性;间作条件下,小麦和蚕豆根系中的IAA含量与根长和根表面积之间存在明显的正相关关系。由此可见,小麦-蚕豆间作能够诱导小麦和蚕豆根系IAA的增加。这种变化可能是驱动间作系统根系形态变化的重要因子。     

局部灼伤对小麦幼苗外源茉莉酸吸收与运转的影响

采用示踪技术探索了3H－JA的运输和分配规律及其受伤害胁迫的影响,外源3H－JA能够在小麦幼苗体内向上和向下运输,局部灼伤其运输与分配都发生了改变,从小麦根系饲喂的3H－JA,在植株内的分布量依序为根＞茎＞叶,时间较长（4h)时分配于心叶的3H0JA大大增加,当叶片受到局部灼伤时3H－JA向地上部的输出量减少,但局部灼伤可加快由心叶饲喂的3H－JA的向下运输,改变3H－JA在小麦幼苗各部位的分配比率。心叶饲喂短时间（5min)时,3H－JA主要积累在受到伤胁迫的展开叶（第2叶）中,向展开叶（第2叶）饲喂 的3H－JA向下运输的速率高于向上运输的速率。推测JA运输及分配的变化可能在植株的防御反应中起重要作用。     

ABA信号转运调节的基因表达与源库动力学分析

通过对拟南芥NCED3、AA03及SDR1蛋白亚细胞定位分析及根系和叶片ABA池的动态库变化研究,结果表明气孔运动的有效ABA信号来自于保卫细胞之外,SDR与ABA前体加工和运输有关。胁迫处理后根系合成酶基因转录水平显著高于叶片,但叶片ABA水平是根系的10倍以上,离体叶片和附体叶片ABA含量测定表明,叶片ABA池的形成主要决定于根源ABA的输入。氟啶酮药剂阻断和遮荫实验说明根系ABA池受叶源类胡萝素前体供应影响。叶片ABA水平受根源ABA和叶源类胡萝素前体库双向转运调节,维管束组织系统可能协同和整合了这一复杂调节机制。该结论为逆境ABA信号转递机制研究和操纵内源ABA含量增强植物抗逆性的应用提供相关资料。     

菌丝桥介入枳根系间抗溃疡病的信号物质响应

对两室根箱试验装置中一分室(供体室)定植的枳Poncirus trifoliata实生苗接种丛枝菌根(AM)真菌Paraglomus occultum,另一分室(受体室)不作任何处理。定植17周后,将溃疡病病菌Xanthomonas axonopodis(Xac)接种于供体植株,研究不同处理受体植株间的信号物质响应差异,揭示菌丝桥的功能。与不接种处理相比,单接种AM真菌、预先接种AM真菌后再接种Xac均显著提高了供体根系一氧化氮(NO)含量,单接种AM真菌只显著提高了受体根系NO含量;单接种Xac显著降低了供体根系脱落酸(ABA)含量,而单接种AM真菌、双接种(AM真菌+Xac)均增加了供体根系ABA含量。双接种处理显著提高了受体ABA含量;与不接种对照相比,接种处理均降低了供体根系玉米素核苷(ZR)含量,同时单接种AM真菌、双接种处理也降低了受体根系ZR含量;AM真菌显著降低了供体和受体根系茉莉酸(JA)含量,但双接种却显著提高了供体和受体根系JA含量;与不接种对照相比,单接种AM真菌或Xac、双接种处理均显著下调供体根系JA合成基因(PtLOX、PtAOS和PtAOC)的表达,单接种Xac显著下调了受体根系PtLOX和Pt AOC的表达,单接种AM真菌显著下调了受体根系PtAOC的表达,而双接种显著上调了受体根系PtAOS基因的表达。表明菌根真菌的菌丝桥传导JA并诱导受体植株中NO、ABA和ZR含量改变以抵御溃疡病菌危害。     

昆虫取食和药剂熏蒸对马尾松针叶脱落酸和茉莉酸含量的影响

脱落酸(ABA)和茉莉酸(JA)在诱导植物产生抗逆性过程中具有重要作用.以马尾松为试验材料,用GC/MS检测了马尾松毛虫咬食、外源茉莉酸甲酯(MeJA)熏蒸和萜烯混合物熏蒸处理4 h后,马尾松损伤针叶、相邻姊妹针叶(处理同一轮未受损伤枝条)、系统上枝针叶(处理枝上一轮枝条)和系统下枝针叶(处理枝下一轮枝条)中ABA和JA含量的变化.结果表明,虫害4 h后,马尾松损伤针叶、相邻姊妹针叶、系统上枝针叶和系统下枝针叶的ABA和JA含量均明显升高.用10 μmol·L^-1 MeJA熏蒸和10 μmol·L^-1萜烯混合物熏蒸健康植株后,各部位枝条针叶中ABA和JA含量明显升高.说明ABA和JA是马尾松苗木伤反应信号传导途径中的重要信号分子,参与了马尾松苗木系统抗性的形成过程.     

异质水分环境下野牛草相连分株间光合同化物的生理整合及其调控

异质环境下,克隆植物通过生理整合机制使资源在分株间实现共享,提高了其对异质性环境的适应能力,具有重要的生态进化意义,研究生理整合机制及其调控机理可为进一步发掘克隆植物应用潜力提供理论依据。以野牛草3个相连分株为材料,对其中一个分株用30%聚乙二醇6000(PEG-6000)模拟水分胁迫,通过Hoagland营养液培养试验,研究了异质水分环境下光合同化物在野牛草相连分株间的生理整合及分株叶片与根系内源激素ABA与IAA含量的变化规律。结果表明,14C-光合同化物在克隆片断内存在双向运输,但以向顶运输为主,异质水分环境下,受胁迫分株光合同化物的输出率明显降低,而与其相邻分株合成的光合同化物向受胁迫分株方向运输率明显增加;异质水分环境下,各分株ABA含量均明显增加,但以受胁迫的分株叶片及根系ABA的含量增加幅度最大,各分株IAA含量较对照均显著下降(P0.05),且以受胁迫分株IAA含量下降幅度最大;各分株叶片与根系ABA/IAA均显著提高(P0.05),相邻分株ABA/IAA增加幅度低于受胁迫分株。异质水分环境影响野牛草克隆分株间光合同化物的生理整合,且ABA与IAA在分株间光合同化物运输与分配过程中具有重要的调节作用。     

JA,ABA在海棠幼苗和枝条中的传输分布以及与水分的关系

采用示踪技术探索了ＪＡ和ＡＢＡ在海棠幼苗和枝条中的传输分布以及与水分的关系,发现ＪＡ的行径主要由韧皮部自上而下运输,ＡＢＡ主要通过木质部由下向上运输。两者均对土壤水分变化敏感,水分亏缺会加速ＪＡ向下运输,促进ＡＢＡ向上运输,推测ＪＡ下运除影响根系生长外,可能与根系对根际环境变化的感应以及根源ＡＢＡ的合成有关。     

外源茉莉酸甲酯诱导抗性与敏感稗草对二氯喹啉酸抗性的差异及机理

以遗传背景一致的抗二氯喹啉酸型和敏感型稗草为供试材料,分析了经外源茉莉酸甲酯(MeJA)预处理后,稗草种群间应对二氯喹啉酸胁迫的抗性水平及4种内源激素含量的变化。结果表明: 外源MeJA处理可显著提高抗性稗草对二氯喹啉酸的抗性,而对敏感稗草的抗性影响不显著。供试稗草间植株体内生长素(IAA)、脱落酸(ABA)、水杨酸(SA)和茉莉酸(JA)的含量及其变化幅度存在显著差异。二氯喹啉酸处理显著增加了稗草ABA、SA和JA的含量,且敏感稗草ABA和JA含量升高幅度明显高于抗性稗草;外源MeJA预处理能显著增强二氯喹啉酸对稗草ABA和JA含量的诱导,且敏感稗草变化幅度较大。表明敏感稗草较快的激素变化不利于其应对除草剂胁迫,抗性稗草相对平缓的激素变化赋予了其对除草剂的适应性,且MeJA处理增强了其对二氯喹啉酸的抗性。稗草植株体内ABA和JA激素信号在其抵抗二氯喹啉酸胁迫过程中可能发挥重要的生物学功能。     

高温胁迫及其信号转导

高温是影响当前农业生产重要的不利环境因素之一。本文综述了4种信号分子脱落酸(ABA),Ca2+,水杨酸(SA),茉莉酸(JA)对高温胁迫的响应以及它们的相互关系,高温胁迫能够诱导ABA,Ca2+,SA的含量升高,并且通过外施ABA,Ca2+,SA,JA都能提高植物的抗热性。作为胞内第二信使,外源Ca2+能够提高植物超氧化物歧化酶(SOD),过氧化氢酶(CAT)以及抗坏血酸氧化酶(APX)的活性,且能提高钙调蛋白水平。ABA诱导的抗热性受胞质游离的Ca2+介导。SA被认为是对胁迫反应所必需的信号分子,H2O2很可能是信号转导链的一部分。JA和ABA在生理功能上有很多相似之处,JA独自或通过提高ABA含量来起作用,JA和SA有不同的生理功能,也有相同的(不过它们的信号转导途径可能不同),最后,提出了今后高温胁迫信号转导研究的一些思路。     

土壤水分亏缺条件下根源信号ABA参与作物气孔调控的数值模拟

建立了根系吸水模型和根源ABA参与作物气孔调控过程相耦合的气孔导度模型,该模型在根源信号ABA的产生项中考虑了根系吸水影响函数和根系密度分布函数.利用该耦合模型模拟大田状况下根源ABA参与玉米气孔行为调控过程,结果表明,由于充分考虑了根区土壤水势和土壤中根长密度分布对根系吸水的影响,较好地反映了土壤不同层次根系吸水强度,更为确切地描述了当土壤水分亏缺时,根系合成ABA的量、各层根系蒸腾流中ABA浓度、木质部ABA浓度以及最终ABA参与对气孔行为的调控作用.     

水分胁迫下ABA由蚕豆根向地上部的运输及其在叶片组织中的分布

蚕豆根装载的３Ｈ－ＡＢＡ可经５．６ｃｍ／ｍｉｎ以上的速率向冠部运输。短时间内（５ｍｉｎ）根运来的ＡＢＡ主要分布在有大量气孔密布的下表皮,但长时间内（３ｈ）则主要分布在对内组织中。抑制蒸腾可降低ＡＢＡ向叶片中的运输积累。光镜放射自显影术显示,根运来的ＡＢＡ可有效地在表皮细胞及保卫细胞的质外体积累。３Ｈ－ＡＢＡ由根向地上部快速运输及其在作用部位的有效积累,说明水分胁迫下蚕豆根部可以通过ＡＢＡ信号的传递控制气孔的行为。     

Root-shoot interactions in mineral nutrition

In this paper four classes of co-operative root-shoot interations are addressed. (I) Nitrogen concentrations in the xylem sap originating from the root and in the phloem sap as exported from source leaves are much lower than those required for growth by apices and developing organs. Enrichment of xylem sap N is achieved by xylem to xylem (X-X) transfer, by which reduced N, but not nitrate, is abstracted from the xylem of leaf traces and loaded into xylem vessels serving the shoot apex. Nitrogen enrichment of phloem sap from source leaves is enacted by transfer of reduced N from xylem to phloem (X-P transfer). Quantitative data for the extent of the contribution of X-X and X-P transfer to the nutrition of young organs of Ricinus communis L. and for their change with time are presented. (II) Shoot and root cooperate in nitrate reduction and assimilation. The partitioning of this process between shoot and root is shifted towards the root under conditions of nitrate- and K-deficiency and under salt stress, while P deficiency shifts nitrate reduction almost totally to the shoot. All four changes in partitioning can be attributed to the need for cation-anion balance during xylem transport and the change in electrical charge occurring with nitrate reduction. (III) Even maintenance of the specificity of ion uptake by the root may – in addition to its need for energy – require a shoot-root interaction. This is shown to be needed in the case of the maintenance of K/Na selectivity under the highly adverse condition of salt stress and absence of K supply from the soil. (IV) Hormonal root to shoot interactions are required in the whole plant for sensing mineral imbalances in the soil. This is shown and addressed for conditions of salt stress and of P deficiency, both of which lead to a strong ABA signalling from root to shoot but result in different patterns of response in the shoot.     

Long Distance Transport of Abscisic Acid in NaCI-Treated Intact Plants of Lupinus albus

Plants of Lupinus albus were grown for 51 d under control (1.1mol m^–3 NaCl) and saline (40 mol m^–3 NaCl) conditions.Plants were harvested and changes of carbon, nitrogen and abscisicacid (ABA) contents of individual organs were determined 41d and 51 d after germination. In the period between the twoharvests xylem and phloem saps were collected and respirationand photosynthesis of individual organs were measured. Usingflows of carbon, C/ABA ratios and increments of ABA flows ofABA in phloem and xylem and rates of biosynthesis and degradationof ABA were calculated. Both under control and saline conditionsnet biosynthesis occurred in the root, the basal strata of leavesand in the inflorescence. Metabolic degradation of ABA tookplace in the stem internodes and apical leaf strata. Salt stress increased xylem transport of ABA up to 10-fold andphloem transport to the root up to 5-fold relative to that ofthe controls. A considerable amount of ABA in the xylem saporiginated from biosynthesis in the roots, i.e. 55% in salt-treatedand smaller than 28% in control plants. The remaining part ofABA in the xylem sap originated from the shoot: it was translocatedin the phloem from fully differentiated leaves towards the rootand from there it was recirculated back to the aerial partsof the plant. The data suggest that ABA may serve as a hormonalstress signal from the root system. Key words: Lupinus albus, salt stress, abscisic acid, long distance transport     

Flows of elements, ions and abscisic acid in Ricinus communis and site of nitrate reduction under potassium limitation.

In a pot experiment Ricinus communis plants were cultivated in quartz sand and supplied daily with a nutrient solution which contained 4 mol m(-3) nitrate as the nitrogen source and either full strength potassium (1.3 mol m(-3), control) or 8% potassium (0.1 mol m(-3), K(+)-limitation). Although the final fresh weight of the whole plant was not affected by K(+)-limitation, the root-shoot ratio was increased due to a relatively increased root growth and inhibited development of younger shoot parts. Owing to K(+)-limitation, photosynthesis was slightly decreased, while dark respiration of the shoot markedly decreased and root respiration was nearly doubled. The transport of carbon in the phloem, and to some extent in the xylem, was greater and the root was favoured in the partitioning of carbon. This was also true for nitrogen and potassium which were both taken up at lower rates, particularly potassium. In these two cases a high remobilization and recycling from the old part of the shoot was observed. By contrast, uptake of sodium was 2.4-fold higher under K(+)-limitation and this resulted in increased flows in the plants, which was discussed generally as a means for charge balance (in combination with a slight increase in uptake of magnesium and calcium). Nitrate reduction took place in the same portion in the root and shoot. This was a shift to the root compared to the control and points to an inhibition of xylem transport caused by limitation of K(+) as an easily permeating countercation. Low K(+) supply also resulted in an increased biosynthesis of ABA in the roots (265%). This caused a slightly increased deposition of ABA in the roots (193%) and a 4.6-fold higher root-to-shoot and a doubled shoot-to-root ABA signal in the xylem or phloem, respectively. The high degradation of ABA in the shoots prevented ABA accumulation there.     

Growth of Zea mays L. plants with their seminal roots only. Effects on plant development, xylem transport, mineral nutrition and the flow and distribution of abscisic acid (ABA) as a possible shoot to root signal

Maize (Zea mays L.) was grown in quartz sand culture eitherwith a normal root system (controls) or with seminal roots only(‘single-rooted’). Development of adventitious rootswas prevented by using plants with an etiolated mesocotyl andthe stem base was positioned 5–8 cm above the sand. Eventhough the roots of the single-rooted plants were sufficientlysupplied with water and nutrients, the leaves experienced waterdeficits and showed decreased transpiration as trans plrationalwater flow was restricted by the constant number of xylem vesselspresent in the mesocotyl. As a consequence of this restriction,transpirational water flow velocities in the metaxylem vesselsreached mean values of 270 m h^–1 and phloem transportvelocities of 5.2 m h^–1. Despite limited xylem transportmineral nutrient concentrations in leaf tissues were not decreasedin single-rooted plants, but shoot and particularly stem developmentwas somewhat inhibited. Due to the lack of adventitious rootsthe shoot:root ratio was strongly increased in the single-rootedplants, but the seminal roots showed compensatory growth comparedto those in control plants. Consistent with decreased leaf conductance,ABA concentrations in leaves of single-rooted plants were elevatedup to 10-fold, but xylem sap ABA concentrations in these plantswere lower than in controls, in good agreement with the well-wateredconditions experienced by the seminal roots. Surprisingly, however,ABA concentrations in tissues of the seminal roots of the single-rooted plants were clearly increased compared to the controls,presumably due to increased ABA import via phloem from the water-stressedleaves. The results are discussed in relation to the role ofABA as a shoot to root signal. Key words: Zea mays, seminal roots, plant development, xylem transport, mineral nutrition, ABA, shoot-to-root signal     

局部灼伤对小麦幼苗外源茉莉酸吸收与运转的影响

采用示踪技术探索了3H-JA的运输和分配规律及其受伤害胁迫的影响.外源3H-JA能够在小麦幼苗体内向上和向下运输,局部灼伤其运输与分配都发生了改变.从小麦根系饲喂的3H-JA,在植株内的分布量依序为根＞茎＞叶,时间较长(4h)时分配于心叶的3H-JA大大增加.当叶片受到局部灼伤时3H-JA向地上部的输出量减少;但局部灼伤可加快由心叶饲喂的3H-JA的向下运输,改变3H-JA在小麦幼苗各部位的分配比率.心叶饲喂短时间(5min)时,3H-JA主要积累在受到伤胁迫的展开叶(第2叶)中.向展开叶(第2叶)饲喂的3H-JA向下运输的速率高于向上运输的速率.推测JA运输及分配的变化可能在植株的防御反应中起重要作用.     

Transport, synthesis and catabolism of abscisic acid (ABA) in intact plants of castor bean (Ricinus communis L.) under phosphate deficiency and moderate salinity

Flows of abscisic acid (ABA) were investigated in whole plantsof castor bean (Ricinus communis) grown in sand culture undereither phosphate deficiency or moderate salinity. Xylem transportof ABA in P-deficient plants was stimulated by a factor of 6whereas phloem transport was affected only very slightly. ABAdeposition into leaves of P-deficient plants was not appreciablydifferent from the controls because of strong net degradationin leaves. Since conjugation of ABA was strongly reduced inall organs of P-deficient plants ABA was presumably metabolizedmainly to phaseic acid and dihydrophaseic acid. The increasedimport of ABA occurred predominantly into fully differentiatedbut not senescent leaves and showed a good correlation withthe inhibition of leaf conductance under P deficiency. As with low-P-plants salt stress increased ABA synthesis inroots and associated transport in the xylem. However, salinitycaused a distinctly greater accumulation of ABA in the leaves,stem segments and the apex than in P-deficient plants. As opposedto P deficiency, ABA export in the phloem from the leaves wasstimulated by salinity. Modelling of ABA flows within an individualleaf over its life cycle showed that young growing leaves importedABA from both phloem and xylem, whereas the adult non-senescentleaves were a source of ABA and thus provided a potential shoot-to-rootstress signal as well as an acceptor for reciprocal signalsfrom root to shoot. In senescing leaves ABA flows and accumulationwere somewhat retarded and ABA was lost in net terms by exportfrom the leaf. Key words: Abscisic acid, phosphorus deficiency, salt stress, phloem and xylem transport     

Foliar application of nitrate or ammonium as sole nitrogen supply in Ricinus communis. II. The flows of cations, chloride and abscisic acid

Following a precultivation with pedospheric nitrogen nutrition, Ricinus plants were supplied with nitrogen solely by spraying nitrate or ammonium solution onto the leaves during the experimental period. The chemical composition of tissues, xylem and phloem exudates was determined and on the basis of the previously determined nitrogen flows (Peuke et al., New Phytologist (1998), 138 , 657–687) the flows of potassium, sodium, magnesium, calcium, chloride and ABA were modelled. These data, which permit quantification of net-uptake, transport in xylem and phloem, and utilization in shoot and root, were compared with results obtained in plants with pedospherically-supplied nitrate or ammonium and data in the literature. Although the overall effects on the chemical composition of supplying ammonium to the leaves were not as pronounced as in pedospherically supplied plants, there were some typical responses of plants fed with ammonium (ammonium syndrome). In particular, in ammonium-sprayed plants uptake and transport of magnesium decreased and chloride uptake was increased compared with nitrate-sprayed plants. Furthermore, acropetal ABA transport in the xylem in ammonium-sprayed Ricinus was threefold higher than in nitrate-sprayed plants. Additionally, concentrations of anions were more or less increased in tissues, particularly in the roots, and transport fluids. The overall signal from ammonium-sprayed leaves without a direct effect of ammonium ions on uptake and transport systems in the root is discussed.     

Is coordination of leaf and root growth mediated by abscisic acid? Opinion

Leaf growth is more inhibited than root growth when the soil is nitrogen-deficient, dry, saline, compacted, or of restricted volume. Similar differential responses in leaf and root growth occur when ABA is applied to plants in well-watered and well-fertilised conditions, and opposite responses are often found in ABA-deficient mutants. ABA levels increase in plants in dry or saline soils, suggesting a regulating role in leaf and root growth in soils of low water potential. In nitrogen-deficient or compacted soils, or soils of restricted volume, ABA only sometimes increases, and in these situations its accumulation may be of secondary importance. Use of ABA-deficient mutants has so far indicated that ABA influences leaf and root growth in unstressed plants, and plants in dry soils, but not in soils that are compacted, of restricted volume, or are nitrogen-deficient.For ABA to determine the relationship between the rate of leaf growth and the rate of root growth, there must be long-distance transport of either ABA itself or a compound that controls ABA synthesis in the growing cells of leaves and roots. ABA invariably increases in xylem sap as the soil becomes dry or saline, and sometimes when it becomes nitrogen-deficient or compacted, however the ABA is of too low a concentration to affect leaf growth. There may be a compound in xylem sap that controls the synthesis of ABA in the leaf, but no such compound has been identified. ABA accumulates in phloem sap of plants in dry or saline soil, but its function in controlling root or leaf growth is unknown.We conclude that ABA affects the ratio of root growth to leaf growth via its independent effects on root and leaf growth, and may regulate the ratio of root to leaf growth via feedforward signals in xylem or phloem, but there is no satisfactory explanation of its mechanism of control.     

Involvement of root ABA and hydraulic conductivity in the control of water relations in wheat plants exposed to increased evaporative demand

We studied the possible involvement of ABA in the control of water relations under conditions of increased evaporative demand. Warming the air by 3°C increased stomatal conductance and raised transpiration rates of hydroponically grown Triticum durum plants while bringing about a temporary loss of relative water content (RWC) and immediate cessation of leaf extension. However, both RWC and extension growth recovered within 30 min although transpiration remained high. The restoration of leaf hydration and growth were enabled by increased root hydraulic conductivity after increasing the air temperature. The use of mercuric chloride (an inhibitor of water channels) to interfere with the rise on root hydraulic conductivity hindered the restoration of extension growth. Air warming increased ABA content in roots and decreased it in shoots. We propose this redistribution of ABA in favour of the roots which increased the root hydraulic conductivity sufficiently to permit rapid recovery of shoot hydration and leaf elongation rates without the involvement of stomatal closure. This proposal is based on known ability of ABA to increase hydraulic conductivity confirmed in these experiments by measuring the effect of exogenous ABA on osmotically driven flow of xylem sap from the roots. Accumulation of root ABA was mainly the outcome of increased export from the shoots. When phloem transport in air-warmed plants was inhibited by cooling the shoot base this prevented ABA enrichment of the roots and favoured an accumulation of ABA in the shoot. As a consequence, stomata closed.