水分胁迫条件下草莓克隆分株间水分调控及其对光合功能的影响

以盆栽草莓(Fragaria×ananassa)为材料研究了水分胁迫下克隆植物草莓母株和子株间的水分调控机制及其与碳同化、光系统Ⅱ激发能分配的关系.实验材料分为匍匐茎连接和剪断两个大组,进行两步实验.第1步实验,对连接组和剪断组的所有母株控水,子株充分供水;4d后进入第2步实验,把连接组分为两小组,对其中一组充分供水子株开始控水,另一组保持不变.结果表明,土壤干旱引起母株叶片失水,并使其净光合速率和气孔导度显著降低.但是连接组中供水良好的子株能有效缓解缺水母株的水分胁迫.当供水良好的子株也开始受到干旱处理的时候,则会加剧与之相连母株的水分胁迫.受胁迫母株可以通过加强渗透调节能力和降低水势从相连子株获取水分.虽然土壤干旱会造成受胁迫母株叶片脱落酸(abscisic acid, ABA)含量的大幅度增加,但是与之相连子株的叶片ABA含量并没有增加;并且气孔导度与ABA变化趋势一致.(1)草莓母株和子株间的水分运输是由二者的水势差驱动的;(2)ABA不会通过匍匐茎在母株和子株间传递并影响相邻子株气孔导度;(3)在水分异质性较大情况下,生理整合可明显提高克隆系统的碳同化能力和光系统Ⅱ激发能利用效率.     

Exploitation of patchy soil water resources by the clonal vine <Emphasis Type="Italic">Ficus tikoua</Emphasis> in karst habitats of southwestern China

The karst habitats of southwestern China are characterized by a highly heterogeneous distribution of water resources. We hypothesized that the clonal integration between connected ramets of the clonal vine Ficus tikoua was an important adaptive strategy to the patchy distribution of water resources in these habitats. We grew ramet pairs (each consisting of a parent and an offspring ramet) in both homogeneously and heterogeneously watered conditions. The offspring ramets were well-watered, whereas their connected parent ramets were randomly assigned to four water treatments: well-watered, mild water stress, moderate water stress, and severe water stress. Increasing water stress decreased leaf water potential, relative water content, net assimilation rate, maximum quantum yield of PSII (F _v/F _m), and biomass of the parent ramets. Subjecting the parents to water stress significantly increased root biomass and root mass ratio (RMR) of their offspring ramets. Exploitation of plentiful water resources through the increased adventitious roots connected to another soil patch permitted the complete restoration of water relations and photosynthetic capacity of offspring ramets after an initial depression. Water relations and gas exchange of the parents were not affected by the water supply to their connected offspring ramets, suggesting that offspring ramets hardly exported water to the stressed parents. However, net assimilation rate and proline content of the offspring ramets increased when they were connected to water-stressed parents. The compensatory photosynthetic responses of offspring ramets connected to stressed parents revealed an increasing trend as the experiment progressed. Morphological and physiological plasticity of F. tikoua in response to heterogeneous water resources allow them to adapt to karst habitats and be suitable candidates for vegetation restoration projects.     

疏叶骆驼刺母株与子株间的水分整合

在未灌溉的土地上, 疏叶骆驼刺(Alhagi sparsifolia)通常不能进行有性繁殖, 克隆繁殖是其种群维持和延续的唯一方式。因此, 克隆性及其相关克隆性状(如水分整合)在疏叶骆驼刺自然种群的维持过程中可能扮演了极其重要的角色。该文通过疏叶骆驼刺母株和子株之间的间隔子切断和给母株补充水分的方法, 研究了母株和子株在各处理下的水势、叶形态和植株生长变化情况。结果表明: (1)间隔子切断后, 疏叶骆驼刺母株和子株正午水势均明显增大(p < 0.01), 说明间隔子切断使得母株和子株水分亏缺值都增大。(2)给母株补水后, 间隔子切断组和间隔子相连组中的母株水势均有明显增加, 同时间隔子相连组的子株水势明显增加(p < 0.01), 而间隔子切断组子株水势没有明显变化(p > 0.05)。(3)间隔子切断组的子株叶片含水率明显低于间隔子相连组子株, 而其株高、冠幅、分枝数和基径的增长量都明显小于间隔子相连组的子株(p < 0.01)。疏叶骆驼刺母株和子株间存在水分整合, 母株会通过根系向子株传输水分。研究成果对塔克拉玛干沙漠南缘的植被恢复以及水资源的合理利用有着重要的意义。     

养分异质条件下结缕草克隆分株生长及光合作用的生理整合

通过对连接和断开的分株进行不同养分处理,研究养分异质条件下结缕草克隆分株生长及光合作用的变化.结果表明:在养分异质条件下,处于中、高养分水平的母株可以提高与其相连子株的叶长、叶宽、根质量、叶质量、光合速率、气孔导度、蒸腾速率和水分利用效率,高养分条件下分别提高16.0%、8.3%、24.4%、58.1%、30.3%、54.0%、9.2%和21.9%,降低根冠比和胞间CO2浓度,在高养分下分别降低21.6%和31.5%;处于中、高养分水平的子株对与其相连母株的生长及光合特性没有显著影响.在养分异质条件下,结缕草母株对子株存在生理整合,养分梯度越大,整合强度越强.结缕草克隆子株可以从母株获益,但母株不能从子株获益,子株是生理整合单向获益者.     

Effects of stolon severing on the expansion of Alternanthera philoxeroides from terrestrial to contaminated aquatic habitats

Few studies have examined the effects of clonal integration (translocation of resources between interconnected ramets) during the expansion of amphibious clonal plants from terrestrial to aquatic habitats. We conducted a greenhouse experiment to simulate the expansion of plants from terrestrial to contaminated aquatic habitats in the amphibious stoloniferous herb Alternanthera philoxeroides (alligator weed). The proximal ramets (i.e. relatively old) of clonal fragments grown in uncontaminated soils were connected to (allowing clonal integration) or disconnected from (preventing clonal integration) distal ramets (i.e. relatively young) grown either in uncontaminated water (control, no CuSO₄) or in four copper‐contaminated water treatments containing 31.25, 62.5, 125 and 250 mg/L CuSO₄, respectively. When a stolon connection was severed, all distal ramets grown in the contaminated water died. When the stolon connection was intact, however, the survival rate of the distal ramets was 85–100% when they were grown at the three lower levels of contamination and 43.75% at the highest level. Moreover, the survival rate and growth of the distal ramets grown in the three lower levels of contamination treatments did not differ from those in the control (uncontaminated water). These results suggest that clonal integration could greatly improve the survival and growth of alligator weed subjected to moderate levels of copper stress. Although clonal integration could also increase the survival rate of the connected distal ramets subjected to the highest level of copper stress (250 mg/L CuSO₄) compared with that of disconnected distal ramets, the survival rate and growth measures were still significantly lower than those in the control. This suggests that clonal integration plays a limited role in the survival and growth of alligator weed when it is subjected to severe stress by high levels of copper contamination.     

Trade-offs among growth, clonal, and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration

The purpose of this article was to study the trade-offs among vegetative growth, clonal, and sexual reproduction in an aquatic invasive weed Spartina alterniflora that experienced different inundation depths and clonal integration. Here, the rhizome connections between mother and daughter ramets were either severed or left intact. Subsequently, these clones were flooded with water levels of 0, 9, and 18 cm above the soil surface. Severing rhizomes decreased growth and clonal reproduction of daughter ramets, and increased those of mother ramets grown in shallow and deep water. The daughter ramets disconnected from mother ramets did not flower, while sexual reproduction of mother ramets was not affected by severing. Clonal integration only benefited the total rhizome length, rhizome biomass, and number of rhizomes of the whole clones in non-inundation conditions. Furthermore, growth and clonal reproduction of mother, daughter ramets, and the whole clone decreased with inundation depth, whereas sexual reproduction of mother ramets and the whole clones increased. We concluded that the trade-offs among growth, clonal, and sexual reproduction of S. alterniflora would be affected by inundation depth, but not by clonal integration.     

Effects of nitrogen addition on clonal integration between mother and daughter ramets of Moso bamboo: a 13C-CO2 pulse labeling study

用¹³C-CO₂脉冲标记法研究氮添加对毛竹母子分株克隆整合的影响 相连分株间的资源共享(即克隆整合)是克隆植物的显著特征。克隆整合使毛竹(Phyllostachys pubescens)对多种环境条件具有较强的适应性。但毛竹通过克隆整合获得性能提升的机制尚不明确。本 研究区分并分析了毛竹光合碳的顶向整合和基向整合,以探究毛竹克隆片段如何在土壤氮异质性条件下 提高整体收益。本研究以由两株不同年龄毛竹分株组成 的克隆片段为研究对象,分株间通过根状茎连接。 每个分株设置氮添加或无氮添加两种处理,并通过单株母株或子株的¹³C-CO₂脉冲标记对光合碳的顶向整合和基向整合进行区分。研究结果表明,子株氮添加显著促进了光合碳由母株向子株的顶向整合,无论母株是否有氮添加,均向氮添加子株转运更多的光合碳。转运自无氮添加母株的光合碳主要分配至氮添加子株的 叶片,而转运自氮添加母株的光合碳则主要分配至根系。与无氮添加处理的子株相比,氮 添 加子株转运更多的光合碳至母株,且该过程不受母株氮添加的影响。转运自氮添加子株的光合碳主要 分配至无氮添加母株的根系和氮添加母株的叶片。这些结果说明,母株优先将更多的资源投入到具有高 养分可利用性的子株,随后子株作为更高效的资源采集点,根据母株的养分状况针对富集资源进行获取。通过这种方式,克隆植物可以降低资源获取成本并提高资源获取效率,最大限度地提升其整体表现。     

Disentangling the contributions of ontogeny and water stress to photosynthetic limitations in almond trees

Very few studies have attempted to disentangle the respective role of ontogeny and water stress on leaf photosynthetic attributes. The relative significance of both effects on photosynthetic attributes has been investigated in leaves of field‐grown almond trees [Prunus dulcis (Mill.) D. A. Webb] during four growth cycles. Leaf ontogeny resulted in enhanced leaf dry weight per unit area (W_a), greater leaf dry‐to‐fresh weight ratio and lower N content per unit of leaf dry weight (N_w). Concomitantly, area‐based maximum carboxylation rate (V_cmax), maximum electron transport rate (J_max), mesophyll conductance to CO₂ diffusion (gm)′ and light‐saturated net photosynthesis (A_max) declined in both well‐watered and water‐stressed almond leaves. Although g_m and stomatal conductance (g_s) seemed to be co‐ordinated, a much stronger coordination in response to ontogeny and prolonged water stress was observed between g_m and the leaf photosynthetic capacity. Under unrestricted water supply, the leaf age‐related decline of A_max was equally driven by diffusional and biochemical limitations. Under restricted soil water availability, A_max was mainly limited by g_s and, to a lesser extent, by photosynthetic capacity and g_m. When both ontogeny and water stress effects were combined, diffusional limitations was the main determinant of photosynthesis limitation, while stomatal and biochemical limitations contributed similarly.     

Physiological integration modifies δ15N in the clonal plant Fragaria vesca,suggesting preferential transport of nitrogen to water-stressed offspring

Background and Aims

One of the most striking attributes of clonal plants is their capacity for physiological integration, which enables movement of essential resources between connected ramets. This study investigated the capacity of physiological integration to buffer differences in resource availability experienced by ramets of the clonal wild strawberry plant, Fragaria vesca. Specifically, a study was made of the responses of connected and severed offspring ramets growing in environments with different water availability conditions (well watered or water stressed) and nitrogen forms (nitrate or ammonium).

Methods

The experimental design consisted of three factors, ‘integration’ (connected, severed) ‘water status’ (well watered, water stressed) and ‘nitrogen form’ (nitrate, ammonium), applied in a pot experiment. The effects of physiological integration were studied by analysing photochemical efficiency, leaf spectral reflectance, photosynthesis and carbon and nitrogen isotope discrimination, the last of which has been neglected in previous studies.

Key Results

Physiological integration buffered the stress caused by water deprivation. As a consequence, survival was improved in water-stressed offspring ramets that remained connected to their parent plants. The nitrogen isotope composition (δ¹⁵N) values in the connected water-stressed ramets were similar to those in ramets in the ammonium treatment; however, δ¹⁵N values in connected well-watered ramets were similar to those in the nitrate treatment. The results also demonstrated the benefit of integration for offspring ramets in terms of photochemical activity and photosynthesis.

Conclusions

This is the first study in which carbon and nitrogen isotopic discrimination has been used to detect physiological integration in clonal plants. The results for nitrogen isotope composition represent the first evidence of preferential transport of a specific form of nitrogen to compensate for stressful conditions experienced by a member clone. Water consumption was lower in plants supplied with ammonium than in plants supplied with nitrate, and therefore preferential transport of ammonium from parents to water-stressed offspring could potentially optimize the water use of the whole clone.     

Photosynthetic response of <Emphasis Type="Italic">Fragaria orientalis</Emphasis> in different water contrast clonal integration

The capacity to exchange resources and non-resource agents is one of the most outstanding features of clonal plants. Contrast between patches in a heterogeneous environment is the main external driving force behind integration effects. It was hypothesized, on the basis of the source–sink hypothesis, that assimilate demand from drought-stressed ramets will result in enhancement of the photosynthesis of well-watered ramets by a mechanism of feedback regulation, that the negative effect of drought on the photosynthesis of drought-stressed ramets will be ameliorated by physiological integration, and that these effects will be enhanced by increasing contrast. A pot experiment was conducted with clonal fragments consisting of two interconnected ramets of Fragaria orientalis. In the experiment, both the connected and the disconnected clonal fragments were divided into three water contrast groups: (1) homogeneous (no contrast) group; (2) low-contrast group; (3) high-contrast group. The photosynthesis and stress tolerance of drought-stressed ramets did not decrease under the support of well-watered ramets when they were connected, allowing clones to maintain their performance in less favorable environments. But the photosynthesis and stress tolerance of drought-stressed ramets decreased with increasing drought-stress when stolons were disconnected. With a feedback regulation process, the photosynthesis of well-watered ramets connected to drought-stressed ramets was enhanced by the latter, which can compensate, at least partially, for the cost of maintaining the stressed ramets. Drought-stressed ramets gained more benefits in a high-contrast environment than in a lower-contrast environment; this can enhance the survival of drought-stressed ramets in unfavorable habitats, especially stressed patches that would otherwise be unexploitable by independent ramets. But photosynthesis of well-watered ramets did not increase with increasing water availability contrast. It can be concluded that photosynthesis and stress tolerance of F. orientalis was affected by clonal integration and by contrasts of water availability.     

Water integration in the clonal emergent hydrophyte, Justicia americana: benefits of acropetal water transfer from mother to daughter ramets

Clonal plant growth is common in aquatic freshwater plants and their success is largely attributed to the sharing of resources (e.g., photosynthates, minerals, and water) within the greater genet. The flow of materials within the clone is largely driven by source-sink dynamics, and in established genets may involve acropetal and/or basipetal flow. During the production of new ramets, however, the mother often provides needed resources to the daughter through acropetal flow. In this study, we consider the role of water sharing in a clonal hydrophyte, Justicia americana, when soil moisture levels around the daughter plant declined. The results of this study indicate that acropetal water sharing is an important function between mother and daughter ramets in this species, as indicated by sustained growth and higher survival in connected daughters residing in water-deprived soils. Interestingly, mother plants, when connected to a drought daughter, began to develop similar xeromorphic features (e.g., greater leaf succulence) even though the parent remained in flooded conditions. We suspect that some physical or chemical signal was conveyed from the daughter to the mother that ??forecast?? water scarcities, which could prepare the entire genet for potential drought conditions.     

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。     

克隆植物沙拐枣的母株和分株对风蚀沙埋的生理生态响应

以中国荒漠区优良的防风固沙克隆灌木沙拐枣为对象,研究了长期风蚀、沙埋环境下沙拐枣母株和克隆分株的同化枝对环境异质性的响应。结果发现:(1)风蚀母株、风蚀分株的叶绿素含量、净光合速率、气孔导度、蒸腾速率、胞间CO2浓度和水分利用效率只有沙埋分株的一半左右,导致同化枝的长度、数量、簇数也仅是沙埋分株的一半,而且风蚀母株的果实宽和果实长也都最小,但浅沙埋有利于沙拐枣的生长和繁殖,表明严峻的风蚀对母株和分株的生长与繁殖都产生了胁迫,但浅沙埋有利于沙拐枣的生长和繁殖。(2)风蚀母株倒伏后同化枝的形态特征是基部优于中部优于顶部,表明严峻风蚀下母株的死亡是从顶部-中部-底部逐渐舍弃的过程。(3)母株的全部根系以及风蚀水平根全部裸露在外但依然能够存活,间接证明沙拐枣克隆整合的方向性——不仅可在分株间进行传递,分株-母株间也可进行传递,否则遭受严峻风蚀胁迫的母株和克隆分株会直接死亡。本研究结果为沙拐枣克隆生长对风沙环境的生态适应机制提供了基础,也是对植物克隆生态学在自然异质环境中研究缺乏的有效补充。     

Physiomorphological response of rose-scented geranium (Pelargonium spp.) to irrigation frequency

The effect of irrigation frequency on leaf physiomorphological processes of rose-scented geranium (Pelargonium capitatum × P. radens cv. Rose) was investigated in a glasshouse study at the Hatfield Experimental Farm of the University of Pretoria, Pretoria, South Africa, from November 2005 to October 2006. Daily, and every 2nd, 3rd, 4th^, and 5th day irrigation were applied as treatments. Leaf samples for electron-microscopic observations were taken one week prior to harvesting, whereafter all plants were re-watered. For each of the irrigation frequency treatments, 50% of the plants were then exposed to a one-week irrigation withholding period (brief stress treatment) prior to harvesting. During this period, physiological properties were recorded on a daily basis to identify or monitor change. Higher irrigation frequency and a brief water stress period increased essential oil yield. Lower irrigation frequency tended to increase the citronellol to geraniol (C:G) ratio to unacceptably high levels (C:G > 3). Upon re-watering, stomatal conductance (G_s) and transpiration rate (R_t) were significantly lower for the lower irrigation frequency treatments, compared to the higher irrigation frequency treatments, while no noticeable differences were observed in water potential (ψ_w) and relative water content (RWC). At the end of the one-week stress period, G_s, R_t, ψ_w and RWC were lower for the plants that were more frequently irrigated compared to the less frequently irrigated treatments. Water stress reduced leaf size, and apparently increased trichome density, whereas the total number of trichomes per leaf remained more or less the same, indicating that total essential oil yield is mainly affected by leaf number (and not by leaf size or trichome number). Stomatal closure was the main water stress avoiding/adaptation mechanism. These results demonstrate that rose-scented geranium plants can make physiomorphological adaptations to save water. However, such a water saving strategy was counter-productive, since it resulted in lower essential oil yield and lower water-use efficiency.     

Movement of Abscisic Acid into the Apoplast in Response to Water Stress in Xanthium strumarium L

The effect of water stress on the redistribution of abcisic acid (ABA) in mature leaves of Xanthium strumarium L. was investigated using a pressure dehydration technique. In both turgid and stressed leaves, the ABA in the xylem exudate, the `apoplastic' ABA, increased before `bulk leaf' stress-induced ABA accumulation began. In the initially turgid leaves, the ABA level remained constant in both the apoplast and the leaf as a whole until wilting symptoms appeared. Following turgor loss, sufficient quantities of ABA moved into the apoplast to stimulate stomatal closure. Thus, the initial increase of apoplastic ABA may be relevant to the rapid stomatal closure seen in stressed leaves before their bulk leaf ABA levels rise.

Following recovery from water stress, elevated levels of ABA remained in the apoplast after the bulk leaf contents had returned to their prestress values. This apoplastic ABA may retard stomatal reopening during the initial recovery period.

     

Clonal integration enhances flood tolerance of Spartina alterniflora daughter ramets

Recently, considerable attention has been paid to the invasion of the clonal plant Spartina alterniflora into coastal wetlands at lower elevations. In this experiment, we tested whether clonal integration improved flood tolerance in S. alterniflora daughter ramets. Daughter ramets at two growth stages (young and old ramets) were flooded to water levels of 0, 9 and 18 cm above the soil surface, and the rhizomes between mother and daughter ramets were either severed or left intact. Biomasses of connected ramets grown in controls or in shallow and deep water treatments were 119%, 108% and 149% higher in the old ramet group than those of severed ramets, respectively, whereas they were 3.0, 3.3 and 11.2 times higher in the young ramet group, respectively. At the end of the experimental period, the shoot height, connected with young ramets, in shallow and deep water treatments increased by 19% and 26%, respectively, over that in the control treatments, whereas the old ramets increased by 11% and 39%, respectively. In contrast, the shoot height of the severed young ramets was 27% and 26% lower in shallow and deep water treatments than in the control treatment, respectively. However, the shoot height of the severed old ramets remained constant with increasing water depth. We conclude that clonal integration enhances the flood tolerance of S. alterniflora daughter ramets, and the trait of clonal integration plays more important roles in severe flooding stress conditions and at early growth stages.     

Water Potential and Component Potentials in Expanded and Unexpanded Leaves of Two Xeric Grasses

Water (ψ), osmotic (ψ_s+ψ_m) and pressure (ψ_p) potentials were measured in three leaf regions of Agropyron dasystachyum and A. smithii grown in the field. Spanner-type thermocouple psychrometers were used to measure ψ and (ψ_m+ψ_m). Absolute water content (AWC) was measured gravimetrically. The ψ and ψ_p were slightly lower in the emerging leaf blade (EBI) than in the last fully emerged leaf blade (FEBI); (ψ_s+ψ_m) and AWC were similar in the two regions. A gradient as large as 0.7 MPa was observed between the EBI and the base of the same emerging leaf (EBs); the latter included the meristematic regions. Although (ψ_s+ψ_m) and ψ_p were generally higher in the EBs, the gradients diminished as the level of stress increased in the shoot. Under moderate water stress the ψ_p of the EBs remained constant relative to the ψ_p in the exposed blades. The large ψ gradient within the growing leaf could have resulted from high resistance imposed by poor vascular development in the intercalary meristem. Ability to maintain a relatively large ψ gradient may be of general significance in buffering the growing region of xeric grass leaves from extreme, short-term fluctuations in water stress that occur in exposed leaf blades.     

High levels of inter-ramet water translocation in two rhizomatous Carex species,as quantified by deuterium labelling

We studied water trnaslocation between interconnected mother and daughter ramets in two rhizomatous Carex species, using a newly developed quantitative method based on deuterium tracing. Under homogeneous conditions, in which both ramets were subjected either to wet or dry soil, little water was exchanged between the ramets. When the ramet pair was exposed to a heterogeneous water supply, water translocation became unidirectional and strongly increased to a level at which 30–60% of the water acquired by the wet ramet was exported towards the dry ramet. The quantity of water translocated was unrelated to the difference in water potential between the ramets, but highly correlated to the difference in leaf area. In both species, the transpiration of the entire plant was similar under heterogeneous and homogeneous wet conditions. This was a direct result of an increase in water uptake by the wet ramet in response to the dry conditions experienced by the interconnected ramet. In C. hirta, the costs and benefits of integration in terms of ramet biomass paralleled the responses of water consumption. This species achieved a similar whole-plant biomass in heterogeneous and homogeneous wet treatments, and water translocation was equally effective in the acropetal and basipetal directions. In C. flacca, responses of biomass and water consumption did not match and, under some conditions, water translocation imposed costs rather than benefits to the plants of this species. It is concluded that enhanced resource acquisition by donor ramets may be of critical importance for the net benefits of physiological integration in clonal plants.     

Water stress‐induced abscisic acid accumulation in relation to reducing agents and sulfhydryl modifiers in maize plant

Signalling process of water stress‐induced abscisic acid (ABA) accumulation was investigated in maize (Zea mays L.) leaf and root tissues. Potent free‐radical scavengers and reducing agents, N‐acetyl cysteine (NAC) and cystein (Cys), significantly inhibited or nearly completely blocked dehydration‐induced ABA accumulation. Dithiothreitol (DTT), a reducing agent but not a free‐radical scavenger, also significantly inhibited such accumulation whereas solely free‐radical scavengers, dimethyl sulphoxide (DMSO) and melatonin (Mela), had no effects. Moreover, water stress‐induced ABA accumulation was not affected either by free radicals, such as superoxide anion and hydrogen peroxide, or by oxidants such as KIO_4. These observations suggest that the blocking of water stress‐induced ABA accumulation resulted from the reducing effect, rather than from anything associated with free radicals. The disulphide bond might be crucial to the reactivity of some signal element(s) in the signalling process of water stress‐induced ABA accumulation. To test the hypothesis, we used a sulfhydryl modifier, iodoacetamide (IOA), and found that it nearly totally blocked the water stress‐induced ABA accumulation. Furthermore, an impermeable sulfhydryl modifier, p‐chloromercuriphenylsulphonic acid (PCMBS), could also inhibit the water stress‐induced ABA accumulation in the leaf tissues. These results indicate that water stress‐perception protein(s) or receptor(s) may be located on the plasmalemma and a sulfhydryl group in the extracellular domain is critical to the reactivity of the speculated water stress receptors. Cys, DTT and IOA did not lead to a decrease of the baseline ABA level, i.e. in non‐stressed roots. Result indicates that their blocking of water stress‐induced ABA accumulation occurred upstream of the ABA biosynthesis pathway, i.e. in the signalling process that initiates such accumulation.     

Does engineering abscisic acid biosynthesis in Nicotiana plumbaginifolia modify stomatal response to drought?

The consequences of manipulating abscisic acid (ABA) biosynthesis rates on stomatal response to drought were analysed in wild‐type, a full‐deficient mutant and four under‐producing transgenic lines of N. plumbaginifolia. The roles of ABA, xylem sap pH and leaf water potential were investigated under four experimental conditions: feeding detached leaves with varying ABA concentration; injecting exogenous ABA into well‐watered plants; and withholding irrigation on pot‐grown plants, either intact or grafted onto tobacco. Changes in ABA synthesis abilities among lines did not affect stomatal sensitivity to ABA concentration in the leaf xylem sap ([ABA]_xyl), as evidenced with exogenous ABA supplies and natural increases of [ABA]_xyl in grafted plants subjected to drought. The ABA‐deficient mutant, which is uncultivable under normal evaporative demand, was grafted onto tobacco stock and then presented the same stomatal response to [ABA]_xyl as wild‐type and other lines. This reinforces the dominant role of ABA in controlling stomatal response to drought in N. plumbaginifolia whereas roles of leaf water potential and xylem sap pH were excluded under all studied conditions. However, when plants were submitted to soil drying onto their own roots, stomatal response to [ABA]_xyl slightly differed among lines. It is suggested, consistently with all the results, that an additional root signal of soil drying modulates stomatal response to [ABA]_xyl.