冰冻胁迫下树木管状分子内腔隙和栓塞的形成及其修复

许多树木管状分子细胞的细胞壁在冰冻期间并不随细胞内的水分迁移到细胞外冰晶上而塌陷。此时细胞内产生负压，负压的产生引起腔隙的形成，腔隙又会引起栓塞，导致树木内水分运输受阻。冻融循环可导致腔隙和栓塞的形成，或者冰冻之后，温度急剧回升时树木组织内的冰晶升华所致。在春季树木的根压得到恢复，从而使腔隙和栓塞部分消除，水分运输又得以畅勇。冰冻胁迫对在高纬度和中高纬度的某些地区的木的生长造成很大的危害，管状分子内腔隙和栓塞的形成就是其中之一，也是引起树木生长衰退或死亡的主要原因。本文对腔隙和栓塞的形成的原因，机理及其恢复进行了综述。     

植物管状细胞栓塞后的重新充注研究进展

在植物吸收水分以后,水分运输对于植物正常的生长发育是非常重要的。在干旱和冬季反复冻融循环以后,植物体内的管状细胞容易充满水蒸气和空气,形成腔 隙和栓塞。腔隙和栓塞的形成对水分在植物体内的运输造成了很大的障碍,从而影响了植物的生长与发育。当植物重新获得水分时,已形成腔隙和栓塞的管状细胞的重新充注能使一部分管状细胞的输水功能得到恢复,从而保证了一些器官的生理功能的正常进行。近些年来,人们对植物管状细胞的重新充注涉及到的许多植物组织和生理过程进行深入的研究,并提出了各种机理。鉴于植物管状细胞形成栓塞后重新充注对植物水分运输的重要生理作用,本文对重新充注的许多机理进行了综合评述。     

植物管状细胞栓塞后的重新充注研究进展

在植物吸收水分以后 ,水分运输对于植物正常的生长发育是非常重要的。在干旱和冬季反复冻融循环以后 ,植物体内的管状细胞容易充满水蒸气和空气 ,形成腔隙和栓塞。腔隙和栓塞的形成对水分在植物体内的运输造成了很大的障碍 ,从而影响了植物的生长与发育。当植物重新获得水分时 ,已形成腔隙和栓塞的管状细胞的重新充注能使一部分管状细胞的输水功能得到恢复 ,从而保证了一些器官的生理功能的正常进行。近些年来 ,人们对植物管状细胞的重新充注涉及到的许多植物组织和生理过程进行深入的研究 ,并提出了各种机理。鉴于植物管状细胞形成栓塞后重新充注对植物水分运输的重要生理作用 ,本文对重新充注的许多机理进行了综合评述     

壳斗科次生木质部水分输导分子间连接通道结构的研究

壳斗科次生木质部水分运输聚合体及其连接通道结构,根据水分运输方向可分纵向和横向两大系统。水分纵向运输通道有导管分子间的穿孔,导管分子侧壁间的管间纹孔,导管分子与无穿孔管状分子间的侧壁纹孔,无穿孔管状分子与无穿孔管状分子间的侧壁纹孔。     

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。     

植物水通道蛋白的运输和调节

水通道蛋白是细胞间和细胞内水分运输的主要通道,其运输和调控对于植物细胞的水分稳态和胁迫响应具有重要作用。本文综述了水通道蛋白运输的分子机制以及结构修饰、门控、膜转运和异源四聚体等调节机制。     

树轮-气候“分异问题”形成机制的研究进展

"分异问题"(divergence problem)最早出现于20世纪90年代。树木年轮学者在阿拉斯加的高山林线研究中,发现树木径向生长对温度敏感性降低,随后在诸多北半球中高纬度地带均有类似报道。这一发现动摇了树轮年代学建立的基本前提假设——"均一性原理",并对基于年轮指数构建数据的科学性提出了挑战。目前,国内外关于"分异问题"形成的原因和机制还存在诸多争论。本文梳理了文献报道的"分异问题"发生的地点、涉及的树种及驱动因子,同时也分析了研究方法等可能引起的误判,以期阐明"分异问题"形成的气候机制。结果表明,"分异问题"主要涉及生长在中高纬度、高海拔地区、对水分和温度要求严格的树种,主要原因是由于气候暖干化加剧土壤水分亏缺(超出水分阈值)或温度升高致使树木生理机能发生改变(超过了正常生长或休眠的温度阈值),导致了树木的气候敏感性降低或者生长衰退。尽管目前关于"分异问题"形成机制的假说还包括:大气CO_2浓度变化、人类活动的影响等,但"分异问题"形成的最可能解释是"阈值假说"。     

林木耗水调控机理研究进展

林木的蒸腾耗水量是造林设计与环境水分研究的重要参数。本文就林木耗水的气孔与非气孔调节机制、木质部空穴和栓塞的发生和恢复机理、树体组织水容等方面进行了综述,对它们在树木水分传输过程中的调控作用和意义开展了探讨。目前在蒸腾气孔调节方面,包括,蒸腾午休、夜间蒸腾、气孔振荡和补偿现象等气孔行为的研究工作有待深入。栓塞木质部和空穴化导管恢复的临界条件与重新充注对植物水分运输的重要生理作用要进一步加强。树体组织水容对树木水分传输和耗水的调控机制问题应加以重视。     

水稻胚性悬浮细胞的包埋脱水法超低温保存

用包埋脱水法冷冻保存水稻胚性悬浮细胞。整个过程包括：胚性悬浮细胞预培养、细胞包埋、二次预培养、包埋细胞脱水、液氮冰冻,细胞解冻和冷冻细胞恢复培养。结果表明,在细胞水分含量为25．17％和蔗糖浓度依次递增以及第2次预培养34d的存活率最好。在培养基中加2．5g·L^-1活性炭有利于细胞的恢复生长。细胞恢复培养后,能再产生愈伤组织,但生长变慢,有约5d的滞后期。     

肝细胞极性与膜蛋白分选的分子机制

肝细胞是高度特化的极性上皮细胞,细胞质膜蛋白的分选和极性转运对于肝细胞极性的建立与维持至关重要.首先,膜蛋白在内质网中合成,随后经高尔基体加工修饰,再由反面高尔基体进一步分选,最后通过膜泡运输等不同的机制分别转运到胆汁腔面或窦状隙面,行使其特殊的功能.近些年来,细胞内负责转运的细胞器和主要的分选信号已逐步被揭示.特别是循环内体也被证明参与了胆汁腔面和窦状隙面膜蛋白的极性分选和转运.肝细胞的极性一旦遭到破坏,将会引起胆汁分泌障碍以及其他肝脏功能的损伤,从而可能导致肝脏糖脂代谢紊乱,甚至丧失正常的生理功能.因此,深入研究肝脏细胞极性的形成与维持机制,将为多种肝脏疾病的预防和治疗寻找到新的方向和靶点,具有重要的理论和临床实践意义.     

Functional and ecological xylem anatomy

Cohesion-tension transport of water is an energetically efficient way to carry large amounts of water from the roots up to the leaves. However, the cohesion-tension mechanism places the xylem water under negative hydrostatic pressure (P_x), rendering it susceptible to cavitation. There are conflicts among the structural requirements for minimizing cavitation on the one hand vs maximizing efficiency of transport and construction on the other. Cavitation by freeze-thaw events is triggered by in situ air bubble formation and is much more likely to occur as conduit diameter increases, creating a direct conflict between conducting efficiency and sensitivity to freezing induced xylem failure. Temperate ring-porous trees and vines with wide diameter conduits tend to have a shorter growing season than conifers and diffuse-porous trees with narrow conduits. Cavitation by water stress occurs by air seeding at interconduit pit membranes. Pit membrane structure is at least partially uncoupled from conduit size, leading to a much less pronounced trade-off between conducting efficiency and cavitation by drought than by freezing. Although wider conduits are generally more susceptible to drought-induced cavitation within an organ, across organs or species this trend is very weak. Different trade-offs become apparent at the level of the pit membranes that interconnect neighbouring conduits. Increasing porosity of pit membranes should enhance conductance but also make conduits more susceptible to air seeding. Increasing the size or number of pit membranes would also enhance conductance, but may weaken the strength of the conduit wall against implosion. The need to avoid conduit collapse under negative pressure creates a significant trade-off between cavitation resistance and xylem construction cost, as revealed by relationships between conduit wall strength, wood density and cavitation pressure. Trade-offs involving cavitation resistance may explain the correlations between wood anatomy, cavitation resistance, and the physiological range of negative pressure experienced by species in their native habitats.     

木本植物木质部的冻融栓塞应对研究进展

冻融栓塞在中高纬度地区木本植物中普遍存在。抗冻融栓塞能力对在寒冷环境中木本植物的生长和安全越冬十分关键, 这直接决定植物分布范围。冻融栓塞是由于冰中气体溶解度低, 木质部水分在低温下冷冻, 使之前水中溶解的气体逸出到导管中, 随后木质部中的冰融化又使气泡扩张而引发的栓塞现象。木质部解剖结构的差异会影响植物的抗冻融栓塞能力, 植物还可以通过调节木质部正压、代谢耗能等方式主动修复冻融栓塞, 也可通过增加树液溶质含量等逃避冷冻, 以减少低温损伤。然而, 与干旱栓塞相比, 目前对木质部冻融栓塞的形成以及植物响应和调节机制的理解不足。为此, 该文首先综述了木质部冻融栓塞的形成机制和植物的逃避、忍耐、修复等3种冻融栓塞的应对策略, 然后总结了木质部抗低温胁迫能力的生理表现、影响因子和评价指标, 并在此基础上讨论了低温抗性、干旱抗性和水力效率之间的多元权衡关系, 最后提出今后该领域中的5个优先研究问题: (1)不同植物冰冻的最低温度阈值; (2)是否存在应对低温胁迫的水力脆弱性分割机制; (3)冻融栓塞修复与代谢消耗的关系; (4)低温抗性、干旱抗性和水力效率之间的权衡关系; (5)抗冻融栓塞性状是否能够纳入经济性状谱系。     

Ultrastructural Evidence That Intracellular Ice Formation and Possibly Cavitation Are the Sources of Freezing Injury in Supercooling Wood Tissue of Cornus florida L

Although cellular injury in some woody plants has been correlated with freezing of supercooled water, there is no direct evidence that intracellular ice formation is responsible for the injury. In this study we tested the hypothesis that injury to xylem ray parenchyma cells in supercooling tissues is caused by intracellular ice formation. The ultrastructure of freezing-stress response in xylem ray parenchyma cells of flowering dogwood (Cornus florida L.) was determined in tissue prepared by freeze substitution. Wood tissue was collected in the winter, spring, and summer of 1992. Specimens were cooled from 0 to -60[deg]C at a rate of 5[deg]C h-1. Freezing stress did not affect the structural organization of wood tissue, but xylem ray parenchyma cells suffered severe injury in the form of intracellular ice crystals. The temperatures at which the ice crystals were first observed depended on the season in which the tissue was collected. Intracellular ice formation was observed at -20, -10, and -5[deg]C in winter, spring, and summer, respectively. Another type of freezing injury was manifested by fragmented protoplasm with indistinguishable plasma membranes and damaged cell ultrastructure but no evidence of intracellular ice. Intracellular cavitation may be a source of freezing injury in xylem ray parenchyma cells of flowering dogwood.     

Freezing of xylem sap without cavitation

Freezing of stem sections and entire twigs of hemlock (Tsuga canadensis) has been demonstrated to occur without increasing the resistance to the movement of water through the frozen part after rewarming. This was interpreted to mean that freezing did not produce cavitation in the xylem sap even though A) the sap was unquestionably frozen; B) it contained dissolved gases; and C) it was under tension before freezing and after. Freezing stem sections of some other evergreen gymnosperms during the summer again produced no evidence for cavitation of the xylem sap. On the other hand, freezing stem sections of some angiosperms invariably increased the resistance to sap flow leading to wilting and death in a few hours when the sap tension was at normal daytime values at the time of freezing. These results were interpreted to mean that the bordered pits on the tracheids of gymnosperms function to isolate the freezing sap in each tracheid so that the expansion of water upon freezing not only eliminates any existing tension but also develops positive pressure in the sap. Dissolved gases frozen out of solution may then be redissolved under this positive pressure as melting occurs. As the bubbles are reduced in size by this ice pressure developed in an isolated tracheid, further pressure is applied by the surface tension of the water against air. If the bubbles are redissolved or are reduced to sufficient small size by the time the tension returns to the sap as the last ice crystals melt, then the internal pressure from surface tension in any existing small bubbles may exceed the hydrostatic tension of the melted sap and the bubbles cannot expand and will continue to dissolve.     

Water transport in trees: current perspectives, new insights and some controversies

This review emphasizes recent developments and controversies related to the uptake, transport and loss of water by trees. Comparisons of the stable isotope composition of soil and xylem water have provided new and sometimes unexpected insights concerning spatial and temporal partitioning of soil water by roots. Passive, hydraulic redistribution of water from moister to drier portions of the soil profile via plant root systems may have a substantial impact on vertical profiles of soil water distribution, partitioning of water within and among species, and on ecosystem water balance. The recent development of a technique for direct measurement of pressure in individual xylem elements of intact, transpiring plants elicited a number of challenges to the century-old cohesion-tension theory. The ongoing debate over mechanisms of long-distance water transport has stimulated an intense interest in the phenomenon and mechanisms of embolism repair. Rather than embolism being essentially irreversible, it now appears that there is a dynamic balance between embolism formation and repair throughout the day and that daily release of water from the xylem via cavitation may serve to stabilize leaf water balance by minimizing the temporal imbalance between water supply and demand. Leaf physiology is closely linked to hydraulic architecture and hydraulic perturbations, but the precise nature of the signals to which stomata respond remains to be elucidated. When water transport in trees is studied at multiple scales from single leaves to the whole organism, considerable functional convergence in regulation of water use among phylogenetically diverse species is revealed.     

Thermodynamic analysis of the interaction of the xylem water and phloem sugar solution and its significance for the cohesion theory

The cohesion theory explains water transport in trees by the evaporation of water in the leaves (transpiration), which in turn generates the tension required for sap ascent, i.e. the flow of pure water from the soil through the root system and the non-living cells of the tree (xylem tracheids) up to the leaves. Only a small part of this water flow entering the leaves is used in photosynthesis to produce sugar solution, which is transported from the leaves through the living cells (phloem) to everywhere in the tree where it is needed and used. The phloem sieves are connected to the xylem tracheids by water transparent membranes, which means that the upflow of pure water and downflow of sugar solution interact with each other, causing the osmotic pressure in the sugar solution (Münch model). In this paper we analyse this interaction with a thermodynamic approach and we show that some open questions in the cohesion theory can then perhaps be better understood. For example, why under a quite high tension the water can flow in the xylem mostly without any notable cavitation, and how the suction force itself depends on the cavitation. Minimizing Gibbs energy of the system of xylem and phloem, we derive extended vapor pressure and osmotic pressure equations, which include gas bubbles in the xylem conduits as well as the cellulose-air-water interface term. With the aid of the vapor pressure equation derived here, we estimate the suction force that the cavitation controlled by the phloem sugar solution can generate at high moisture contents. We also estimate the suction force that the transpiration can generate by moisture gradient at low moisture contents. From the general osmotic pressure equation we derive an equation for calculating the degree of cavitation with different sugar solution concentrations and we show the conditions under which the cavitation in the xylem is totally avoided. Using recent field measurement results for a Scotch pine, the theory is demonstrated by showing its predictions for possible amounts of cavitation or embolism from morning hours to late afternoon.     

The role of freezing in setting the latitudinal limits of mangrove forests

Mangrove trees dominate coastal vegetation in tropical regions, but are completely replaced by herbaceous salt marshes at latitudes above 32 degrees N and 40 degrees S. Because water deficit can increase damage caused by freezing, we hypothesized that mangroves, which experience large deficits as a result of saline substrates, would suffer freeze-induced xylem failure. Vulnerability to freeze-induced xylem embolism was examined in the most poleward mangrove species in North America, in an area where freezing is rare but severe, and in Australia, in an area where freezing is frequent but mild. Percentage loss in hydraulic conductivity was measured following manipulations of xylem tension; xylem sap ion concentration was determined using X-ray microanalysis. Species with wider vessels suffered 60-100% loss of hydraulic conductivity after freezing and thawing under tension, while species with narrower vessels lost as little as 13-40% of conductivity. These results indicate that freeze-induced embolism may play a role in setting the latitudinal limits of distribution in mangroves, either through massive embolism following freezing, or through constraints on water transport as a result of vessel size.