维管植物木质部水分传输过程的影响因素及研究进展

维管植物的水分传输过程一直是植物学家关注的重点,木质部是植物体内长距离运输的组织之一,为维管植物提供了一个将水分从根运输到叶的低阻力通道。木质部在结构上形成了一个相互连通的网络管道结构,外部环境要素的改变(水分胁迫、氮沉降、光照等)引起的植物体内水势及木质部网络管道结构的变化直接影响植物的水分传输过程。本文从植物水分传输驱动力和内部木质部微观结构出发,总结了水分传输机理过程,概述了影响木质部水分传输的直接和间接因素,并在全球变化背景下,在植物内部观测技术、考虑多因子的综合作用等研究方面进行了展望。     

植物木质部导管栓塞

植物木质部栓塞直接影响植物体内的水分传输,文章对近年来植物木质部导管栓塞的时空分布规律、栓塞修复的微观过程,以及根压与作物木质部导管栓塞的关系研究进展作了概述。     

植物根-土界面水分再分配研究方法与影响因素

水分再分配是根-土界面根系对水分在垂直或水平方向的输导过程,对水分再分配的研究最早是在实验室盆栽条件下进行的,而盆栽测定的最基本方法就是分根法。实验室条件下,土壤水分的测定包括γ射线密度法、称量法、烘干法和时域反射仪（TDR）等。野外条件下,对根-土界面水分再分配的研究基本上有4种方法,利用热电隅干湿球温度计测定土壤水势的昼夜及季节波动、氢同位素示踪法、根木质部液流的测量和TDR测定土壤体积含水量。影响水分再分配的因素主要有土壤水分、土壤质地、植物种类及蒸腾作用等。在今后的研究中,应当加强对植物根-土界面水分再分配发生条件的研究,明确各影响因素与水分再分配发生的频率和数量的关系,从而为更好地利用其水分调节功能提供理论基础。     

用热脉冲速度记录仪（HPVR）测定树干液流

树木蒸腾耗水是环境生态平衡(水分)的重要因素。由于树体高大,环境、时间、空间变异因素复杂,测定工作十分困难。在林木生态系统中,水分运动的途径是,树木根部吸收土壤水分,通过树干(木质部上升液流)输送到树冠部,从叶表面蒸腾散失到大气中,即所谓“土壤-植物-大气连续系统”。在此过程中,树干是水流通道的咽喉部位,树干液流量的大小制约着冠部蒸腾量的变化。因此,可以用测定树干部液流的方法确定树冠的蒸腾耗水     

树木树液上升机理研究进展

水分在植物体内的运输一直是很多植物生理生态学家所关注的一个重要问题。介绍了内聚力学说的基本假设和其存在争议,总结了近年来这一研究领域的几个热点问题,主要包括：（1）木质部栓塞及其恢复机理;（2）木质部压力探针和压力室法测定的木质部张力值不一致的现象及其可能原因;（3）补偿压学说;（4）不同界面层张力以及输水管道的毛细作用力、薄壁细胞膨压和木质部渗透压、逆向蒸腾等在树木汁液上升中的贡献;（5）最近发现的存在于木质部导管伴胞和韧皮部薄壁细胞等质膜中的水孔蛋白在植物水分运输中的调控作用等。这些方面在解释树木的树液上升中都起着重要的作用。     

不同冲洗液对毛白杨和油松枝条水力导度和抵抗空穴化能力测定值的影响

植物通过木质部管道系统进行水分运输, 木质部的水分运输效率和抗空穴化能力等水力结构特征对于植物物种的分布、抗逆能力等方面起关键性作用。目前, 国内外学者一般采用“冲洗法”进行木质部水力结构研究, 然而在该方法中使用的不同冲洗溶质可能对植物木质部水力结构等产生较大影响, 因此该文研究了3种溶质的冲洗溶液对毛白杨(Populus tomentosa)和油松(Pinus tabulaeformis)枝条的水力导度和抵抗空穴化能力的影响。实验结果表明: 相对于去离子水, 用0.01 mol·L^-1的草酸和0.03 mol·L^-1KCl溶液作为冲洗溶液, 均导致毛白杨木质部导管和油松管胞的水力导度测定值的增大。KCl导致毛白杨和油松木质部抵抗空穴化能力测定值的提高, 草酸导致杨树抵抗空穴化能力测定值增强, 但导致油松抗空穴化能力显著(p<0.01)减弱。小枝水平上, 毛白杨和油松的水分运输效率和抗空穴化能力之间没有显著相关性。另外, 在截枝实验中发现, 毛白杨小枝木质部水力导度随长度增加变化不大, 而油松枝条的木质部水力导度有逐渐增大的趋势。以上的实验结果表明不同溶质下毛白杨和油松枝条的木质部水力导度和抵抗空穴化能力不同, 草酸和KCl可能对木质部管道系统及纹孔处的果胶等产生作用, 从而使毛白杨和油松的水力结构发生变化。毛白杨与油松水力结构在去离子水、草酸和KCl的作用下的不同结果及两物种截枝试验下水力导度的不同变化趋势表明, 导管运输系统和管胞运输系统可能具有不同的水分运输影响因素。     

根压与根系的主动吸水

植物根系吸收水分有两种类型:被动吸水和主动吸水,被动吸水的动力是蒸腾拉力,主动吸水的动力是根压。我认为根压不是植物根系吸水的动力,而应当是植物根系吸水的结果。理由有三点: 第一,根压的定义没有“根压是根系主动吸水的动力”的含义。“根系的生理活功使液流上升的压力称为根压”。“根压是由于根系代谢活动的结果,使木质部导管分子内产生     

植物根系提水作用研究述评

根系提水作用是植物在干旱生境下,通过根系将深层湿润土壤中的水分提升至浅层干燥土壤中的一种生理过程,不同植物具有不同的根系提水强度。这一研究的基本方式是分根法,而时域反射仪(TDR)法和中子水分仪法则是近年来较受重视的测定方法,并逐步由室内测定向大田测定过渡。研究的主要内容可归纳为五个方面,其中在根系提水作用与植物蒸腾作用、浅层土壤营养的有效性以及与植物遗传性的关系等方面,研究结果较为一致,但在提水的数量和发生提水作用时土壤水分状况等方面,研究结果不尽相同;对植物光合、呼吸等其他代谢的影响以及室内测定结果与大田条件一致性等问题的研究,报道较少,特别是作用机理的研究。因此,深入这方面的研究对于进一步提高植物水分利用率,促进旱地节水农业和畜牧业的发展具有重要的意义。     

盐胁迫下大麦根系木质部压力的自调节现象

用植物木质部压力探针测定的结果表明,水培大麦幼苗根的木质部压力在环境条件恒定不变时始终保持波动,并且在受到轻度的盐胁迫和当盐胁迫解除时表现出高度的自调节现象.这种波动和自调节现象将对植物水势的测定和根的径向反射系数的测定产生很大的影响,并可能与植物的抗盐性有关.小麦根在同样条件下未表现出上述现象.     

桂西南喀斯特季雨林木本植物的水力安全

桂西南喀斯特地区生物多样性丰富、特有种多,同时也是石漠化问题较为严重的区域。由于该喀斯特地区土层浅薄、岩石裸露、表层储水能力差,植物在干旱季节经常会受到水分胁迫。植物水力学特征不仅是探讨喀斯特地区植物的生理生态适应性的关键,还能够为石漠化地区的植被恢复提供重要参考。该研究测定了桂西南喀斯特季雨林17种代表性木本植物(包括不同生活型、叶片习性和生境)的木质部脆弱性曲线、最低水势、叶片膨压丧失点和边材密度等水力性状,结果发现:(1)喀斯特植物木质部导水率丧失50%时的水势值(P50)的种间差异较大(–0.51––2.51MPa),其中常绿种的抗栓塞能力比落叶种强;(2)喀斯特植物的木质部水力安全边界值(最低水势与P50之间的差值)的均值为0.36 MPa,说明喀斯特森林植物在自然最低水势状况下木质部发生栓塞的程度较高;但是不同植物种间存在显著差异,这可能与喀斯特峰丛洼地生境的复杂性以及物种不同的抗旱策略有关;(3)由于喀斯特植物水分适应机制的多样化,导致木质部水力安全边界与叶片膨压丧失点、边材密度的相关性并不显著。在区域气候干热化的背景下,结合喀斯特植物的栓塞脆弱性和长期水势监测(尤其极端干旱事件)分析它们的水力安全,对预测未来喀斯特森林物种分布和群落动态具有重要的指示作用。     

During measurements of root hydraulics with pressure probes, the contribution of unstirred layers is minimized in the pressure relaxation mode: comparison with pressure clamp and high-pressure flowmeter

The effects of unstirred layers (USLs) at the endodermis of roots of young maize plants (Zea mays L.) were quantified, when measuring the water permeability of roots using a root pressure probe (RPP) in the pressure relaxation (PR) and pressure clamp (PC) modes. Different from PRs, PCs were performed by applying a constant pressure for certain periods of time. Experimental data were compared with results from simulations based on a convection versus diffusion (C/D) model, with the endodermis being the main barrier for solutes and water. Solute profiles in the stele were calculated as they occurred during rapid water flows across the root. The model quantitatively predicted the experimental finding of two distinct phases during PRs, in terms of a build-up of concentration profiles in the stele between endodermis and xylem vessels. It also predicted that, following a PC, half-times (T1/2) of PRs increased as the time used for clamping (and the build-up of USLs) increased. Following PCs of durations of 15, 30 and 60 s, T1/2 increased by factors of between 2.5 and 7.0, and water permeability of roots (root hydraulic conductivity, Lpr) was reduced by the same factors. When root pressure was immediately taken back to the original equilibrium root pressure following a PC, there was a transient uptake of water into the root stele (transient increase of root pressure), and the size of transients rose with time of clamping, as predicted by the model. The results indicated that the 'real' hydraulic conductivity of roots should be measured during initial water flows, such as during the rapid phase of PRs, when the effect of USLs was minimized. It was discussed that 'pressure-propagation effects' could not explain the finding of two phases during PRs. The results of USL effects threw some doubt on the use of PC and high-pressure flowmeter (HPFM) techniques with roots, where rigorous estimates of USLs were still missing despite the fact that large quantities of water were forced across the root.     

Water and solute flow through mung bean roots under applied pressure

The technique of applying hydrostatic pressure on the root medium to study water and solute flows through excised plant roots and to study various characteristics of roots in relation to flow has been used by many workers but flows in excised roots have not been compared with those in intact transpiring plants. In the present study this comparison has been made using mung bean roots. Results show that excised roots under pressure lack the ion selectivity which is observed in intact plant roots and conduct salt many times higher than salt flows through intact plant roots. The role of stem resistance in the rates of water and salt flow through roots has been discussed. The suitability of this technique for solute flow studies through mung bean roots is questioned.     

Xylem Flow and its Driving Forces in a Tropical Liana: Concomitant Flow-Sensitive NMR Imaging and Pressure Probe Measurements

Abstract: Flow-sensitive NMR imaging and pressure probe techniques were used for measuring xylem water flow and its driving forces (i.e., xylem pressure as well as cell turgor and osmotic pressure gradients) in a tropical liana, Epipremnum aureum. Selection of tall specimens allowed continuous and simultaneous measurements of all parameters at various distances from the root under diurnally changing environmental conditions. Well hydrated plants exhibited exactly linearly correlated dynamic changes in xylem tension and flow velocity. Concomitant multiple-probe insertions along the plant shoot revealed xylem and turgor pressure gradients with changing magnitudes due to environmental changes and plant orientation (upright, apex-down, or horizontal). The data suggest that in upright and - to a lesser extent - in horizontal plants the transpirational water loss by the cells towards the apex during the day is not fully compensated by water uptake through the night. Thus, longitudinal cellular osmotic pressure gradients exist. Due to the tight hydraulic coupling of the xylem and the tissue cells these gradients represent (besides the transpiration-induced tension in the xylem) an additional tension component for anti-gravitational water movement from the roots through the vessels to the apex.     

Molecular mechanisms of attachment of Rhizobium bacteria to plant roots

Attachment of bacteria to plant cells is one of the earliest steps in many plant-bacterium interactions. This review covers the current knowledge on one of the best-studied examples of bacterium-plant attachment, namely the molecular mechanism by which Rhizobium bacteria adhere to plant roots. Despite differences in several studies with regard to growth conditions of bacteria and plants and to methods used for measuring attachment, an overall consensus can be drawn from the available data. Rhizobial attachment to plant root hairs appears to be a two-step process. A bacterial Ca(2+)-binding protein, designated as rhicadhesin, is involved in direct attachment of bacteria to the surface of the root hair cell. Besides this step, there is another step which results mainly in accumulation and anchoring of the bacteria to the surface of the root hair. This leads to so-called firm attachment. Depending on the growth conditions of the bacteria, the latter step is mediated by plant lectins and/or by bacterial appendages such as cellulose fibrils and fimbriae. The possible role of these adhesions in root nodule formation is discussed.     

Applications of the compensating pressure theory of water transport

Some predictions of the recently proposed theory of long-distance water transport in plants (the Compensating Pressure Theory) have been verified experimentally in sunflower leaves. The xylem sap cavitates early in the day under quite small water stress, and the compensating pressure P (applied as the tissue pressure of turgid cells) pushes water into embolized vessels, refilling them during active transpiration. The water potential, as measured by the pressure chamber or psychrometer, is not a measure of the pressure in the xylem, but (as predicted by the theory) a measure of the compensating pressure P. As transpiration increases, P is increased to provide more rapid embolism repair. In many leaf petioles this increase in P is achieved by the hydrolysis of starch in the starch sheath to soluble sugars. At night P falls as starch is reformed. A hypothesis is proposed to explain these observations by pressure-driven reverse osmosis of water from the ground parenchyma of the petiole. Similar processes occur in roots and are manifested as root pressure. The theory requires a pump to transfer water from the soil into the root xylem. A mechanism is proposed by which this pump may function, in which the endodermis acts as a one-way valve and a pressure-confining barrier. Rays and xylem parenchyma of wood act like the xylem parenchyma of petioles and roots to repair embolisms in trees. The postulated root pump permits a re-appraisal of the work done by evaporation during transpiration, leading to the proposal that in tall trees there is no hydrostatic gradient to be overcome in lifting water. Some published observations are re-interpreted in terms of the theory: doubt is cast on the validity of measurements of hydraulic conductance of wood; vulnerability curves are found not to measure the cavitation threshold of water in the xylem, but the osmotic pressure of the xylem parenchyma; if measures of xylem pressure and of hydraulic conductance are both suspect, the accepted view of the hydraulic architecture of trees needs drastic revision; observations that xylem feeding insects feed faster as the water potential becomes more negative are in accord with the theory; tyloses, which have been shown to form in vessels especially vulnerable to cavitation, are seen as necessary for the maintenance of P, and to conserve the supplementary refilling water. Far from being a metastable system on the edge of disaster, the water transport system of the xylem is ultrastable: robust and self-sustaining in response to many kinds of stress.     

Water Pumping Activity of the Root System in the Process of Cross-Adaptation of Sunflower Plants to Hyperthermia and Water Deficiency

To elucidate the mechanisms of cross-adaptation, we investigated the effect of heat shock (HS, for 2 h at 45°C) on leaf tolerance to overheating and exudation by roots detached from 25–30-day old sunflower (Helianthus annuus L.) plants. It was demonstrated that preheating enhanced considerably leaf tolerance and activated root exudation, especially under water deficiency produced by plant transfer to the hypertonic NaCl solution (17 mM). Under water deficiency conditions, the roots of HS-treated plants pumped water against the osmotic pressure (OP) gradient between the exudate and the external solution. Therefore, we concluded that this pumping was realized due to a metabolic (non-osmotic) constituent of root pressure. In the roots of plants that were not treated with HS, the OP gradient became positive. This fact implies that the HS-pretreatment of plants retarded the penetration of sodium and chlorine ions into roots. The data obtained demonstrate that HS induced a cross-adaptation of plants to high temperature and water deficiency. Such cross-adaptation involves, as an important component, an acceleration of water metabolism, including an enhanced water pumping activity of root system.     

Comparison between gradient-dependent hydraulic conductivities of roots using the root pressure probe: the role of pressure propagations and implications for the relative roles of parallel radial pathways

Hydrostatic pressure relaxations with the root pressure probe are commonly used for measuring the hydraulic conductivity (Lp(r)) of roots. We compared the Lp(r) of roots from species with different root hydraulic properties (Lupinus angustifolius L. 'Merrit', Lupinus luteus L. 'Wodjil', Triticum aestivum L. 'Kulin' and Zea mays L. 'Pacific DK 477') using pressure relaxations, a pressure clamp and osmotic gradients to induce water flow across the root. Only the pressure clamp measures water flow under steady-state conditions. Lp(r) determined by pressure relaxations was two- to threefold greater than Lp(r) from pressure clamps and was independent of the direction of water flow. Lp(r) (pressure clamp) was two- to fourfold higher than for Lp(r) (osmotic) for all species except Triticum aestivum where Lp(r) (pressure clamp) and Lp(r) (osmotic) were not significantly different. A novel technique was developed to measure the propagation of pressure through roots to investigate the cause of the differences in Lp(r). Root segments were connected between two pressure probes so that when root pressure (P(r)) was manipulated by one probe, the other probe recorded changes in P(r). Pressure relaxations did not induce the expected kinetics in pressure in the probe at the other end of the root when axial hydraulic conductance, and probe and root capacitances were accounted for. An electric circuit model of the root was constructed that included an additional capacitance in the root loaded by a series of resistances. This accounted for the double exponential kinetics for intact roots in pressure relaxation experiments as well as the reduced response observed with the double probe experiments. Although there were potential errors with all the techniques, we considered that the measurement of Lp(r) using the pressure clamp was the most unambiguous for small pressure changes, and provided that sufficient time was allowed for pressure propagation through the root. The differences in Lp(r) from different methods of measurement have implications for the models describing water transport through roots and the potential role of aquaporins.     

Stomatal responsiveness to leaf water status in common bean (Phaseolus vulgaris L.) is a function of time of day

Stomatal response to leaf water status was experimentally manipulated by pressurizing the soil and roots of potted common bean plants enclosed in a custom‐built root pressure chamber. Gas exchange was monitored using a whole‐plant cuvette and plant water status using in situ leaf psychrometry. Bean plants re‐opened their stomata upon pressurization, but the extent of re‐opening was strongly dependent on the time of day when the soil was pressurized, with maximum re‐opening in the morning hours and limited re‐opening in the afternoon. Neither leaf nor xylem abscisic acid concentrations could explain the reduced response to pressurization in the afternoon. The significance of this phenomenon is discussed in the context of circadian rhythms and of other recent findings on the ‘apparent feed‐forward response’ of the stomata of some species to vapour pressure deficit.     

Unusual stomatal behaviour on partial root excision in wheat seedlings

The excision of four out of five primary roots of wheat (Triticum durum Desf.) seedlings often leads to an enhanced rate of transpiration. Surprisingly this enhancement could be maintained for several hours after root excision and was particularly likely to occur at low irradiances or high atmospheric humidity. This long‐term enhancement could not be explained in terms of conventional hydropassive stomatal effects. Elevated rates of transpiration were associated with and possibly caused by increased cytokinin concentrations in shoots of plants with partially excised roots. The single root remaining after excision was able to maintain an adequate water uptake for the continued enhanced transpiration, after only a short transient reduction in leaf water content. The enhanced capacity for water uptake by the remaining root was confirmed by measuring the water flow from detached roots at negative hydrostatic pressure. Even without additional suction, flow from the reduced root system increased about 1.5 h after the start of treatment, suggesting an increase in membrane permeability for water. Although abscisic acid (ABA) concentrations in the roots increased after the root excision treatment, there was no evidence for any enhanced concentration in the xylem sap. The possible role that this accumulation of ABA in roots may have in the apparent increase in hydraulic conductivity after root excision is discussed.