胞间连丝与大分子物质的胞间转运机制

胞间连丝是相邻细胞间共质体运输的桥梁。基于对胞间连丝分子组成及超微结构的研究,不同学者提出了不同的胞间连丝结构模型。对其功能的研究表明,胞间连丝在物质运输、信息传递、病毒的周身感染等方面都具有重要作用。文章就胞间连丝的结构、分子组成及病毒介导的大分子胞间转移,以及对内源蛋白质的胞间转运机制诸方面的研究进展作了概述。     

胞间连丝研究的进展

胞间连丝为多细胞植物有机体提供了一个直接的细胞间物质运输和信息传递的细胞质通道,把一个个独立的“细胞王国”转变成相互连接的共质体,它是当今细胞生物学中十分活跃的研究领域。日益增多的研究结果揭示,胞间连丝协调基因表达和许多的细胞生理生化过程,对细胞的分裂与分化、形态发生、植物体的生长与发育,以及植物对环境的反应与适应等诸方面都起着十分重要的作用。本文仅就胞间连丝结构的多样性;胞间通道的调节因子;大分子蛋白质和核酸的胞间运输;胞间连丝阻断和共质体分区的形成及其与形态发生、休眠和抗逆性的关系等几个方面的新进展做一个简要的综述,借此例证胞间连丝在植物生命活动中的重要意义。     

胞间连丝研究的进展

胞间连丝为多细胞植物有机体提供了一个直接的细胞间物质运输和信息传递的细胞质通道,把一个个独立的“细胞王国”转变成相互连接的共质体,它是当今细胞生物学中十分活跃的研究领域。日益增多的研究结果揭示,胞间连丝协调基因表达和许多的细胞生理生化过程,对细胞的分裂与分化、形态发生、植物体的生长与发育,以及植物对环境的反应与适应等诸方面都起着十分重要的作用。本文仅就胞间连丝结构的多样性;胞间通道的调节因子;大分子蛋白质和核酸的胞间运输;胞间连丝阻断和共质体分区的形成及其与形态发生、休眠和抗逆性的关系等几个方面的新进展做一个简要的综述,借此例证胞间连丝在植物生命活动中的重要意义。     

胞间连丝与大分子物质的胞间转移

胞间连丝是细胞间细胞器,是细胞间通讯的直接途径。一般认为,胞间连丝允许通过物质的分子量上限(SEL)是800～1000 Da．近年来研究的许多证据表明,胞间连丝的SEL随组织种类及其生理状况而异。在某些情况下,它可以允许大分子物质通过,如病毒运动蛋白与胞间连丝相互作用,使病毒通过胞间连丝转移。玉米突变体 kn1基因异常表达的KN1可使包括表皮在内的各层组织结瘤,KN1是细胞间移动的信息物,P-蛋白可由伴胞通过胞间连丝转移到筛管。某些组织中胞间连丝很高的SEL和发育过程胞间连丝SEL的变化可能在植物发育调控中有重要作用。本文对大分子通过胞间连丝转移的机理进行了讨论。     

胞间连丝与大分子物质的胞间转移

胞间连丝是细胞间细胞器,是细胞间通讯的直接途径。一般认为,胞间连丝允许通过物质的分子量上限（ＳＥＬ）是８００～１０００Ｄａ．近年来研究的许多证据表明,胞间连丝的ＳＥＬ随组织种类及其生理状况而异。在某些情况下,它可以允许大分子物质通过,如病毒运动蛋白与胞间连丝相互作用,使病毒通过胞间连丝转移。玉米突变体ｋｎ１基因异常表达的ＫＮ１可使包括表皮在内的各层组织结瘤,ＫＮ１是细胞间移动的信息物,Ｐ蛋白可由伴胞通过胞间连丝转移到筛管。某些组织中胞间连丝很高的ＳＥＬ和发育过程胞间连丝ＳＥＬ的变化可能在植物发育调控中有重要作用。本文对大分子通过胞间连丝转移的机理进行了讨论。     

植物病毒长距离转运的分子机理

植物病毒侵入寄主细胞后,其局部侵染和系统侵染的形成涉及病毒在植物体内二种不同的转运模式:经过叶肉细胞胞间连丝来实现的胞间转运(cel-to-cell movement)和经过植物维管系统的韧皮部筛管来实现的长距离转运(long-distance transport)[1].近十年来对胞间转运的大量研究,尤其是对TMV在烟草叶肉细胞间转运机理的出色研究,使人们逐步明晰了病毒胞间转运的一些基本步骤及转运机理,建立起了植物病毒胞间转运机理研究的基本模式[2-5].与此同时,因病毒的长距离转运是其实现系统侵染的关键过程,人们对病毒长距离转运机理的研究也积累了相当多的工作,该方面的研究日益成为植物病毒学研究的一个重要内容.本文拟对病毒长距离转运过程中所涉及的病毒因子、病毒-寄主的互作及病毒进出韧皮部筛分子的可能方式作一概述.     

植物胞间连丝的形成和结构变化

通过对胞间连丝的起源,形成及其调控物质运输机理的讨论,说明胞间连丝是物质运输和信息传递的重要通道。并且比较了高等植物和低等植物胞间连线的发生和次生变化。由于酶的作用,胞间连丝向胞间通道转化,形成了约１００￣１０００ｎｍ的开放性通道。     

植物胞间连丝的形成和结构变化

 通过对胞间连丝的起源、形成及其调控物质运输机理的讨论,说明胞间连丝是物质运输和信息传递的重要通道。并且比较了高等植物和低等植物胞间连丝的发生和次生变化。由于酶的作用,胞间连丝向胞间通道转化,形成了约100~1000nm的开放性通道。     

植物病毒在细胞间转运的机理探讨

植物病毒在寄主体内的移动包括细胞间转运和系统性转运两个部分。在这两个过程中,如何有效地利用和修饰胞间连丝,是病毒成功侵染的关键。病毒通过编码运动蛋白与寄主因子互作靶定于细胞质膜,然后通过一系列复杂机制修饰胞间连丝从而顺利完成细胞间转运。综述了植物病毒在细胞间转运过程中与寄主发生的一系列互作,着重阐述了病毒与胞间连丝之间互作的机制,旨在为相关研究工作提供参考。     

胞间连丝的次生形成和次生变化

本文介绍了胞间连丝次生形成和次生变化的研究进展。用统计特定细胞壁区段上胞间连丝数量与密度的变化,电镜观察嫁接组合中接穗与砧水间细胞壁上胞间连丝的形成等方法,证明了在植物生长发育过程中,存在着胞间连丝的次生形成。在某些特定部位,某一发育阶段,已形成的胞间连丝常会发生可逆的次生变化,这种变化和植物发育过程中的共质体隔离以及物质运输的调节有关。     

蒜鳞片薄壁细胞衰败过程中类外连丝的形成及其共质传输能力的调节（简报）

在植物生长发育过程中,由于局部组织（细胞）的衰退或脱落．存活细胞与衰亡（或脱落）细胞分界壁上的胞间连丝衍变为类外连丝结构。这已先后在小麦胚胎发育、幼胚分化、蒜鳞茎休眠进程中以及玉米根冠组织中得到了论证。前文已曾报道．在玉米根冠细胞生长脱落进程中,脱落细胞与相邻细胞间胞间连丝有被拉伸、断裂形成类外连丝的结构变化．并经药理学试验表明。肌动蛋白和肌球蛋白共同参与了类外连丝通透性的调节。     

蒜鳞片休眠进程中胞间联络的变化及类外连丝结构与功能的研究

利用电子显微镜术,对蒜休眠进程中鳞片薄壁细胞间胞间联络的特征进行了实验观察,发现不同时期胞间联络具有随细胞间生理关系密切程度而呈现相应结构变化的特点。并观察到萌芽期鳞片中衰败细胞与存活细胞之间有类外连丝型胞间连丝的存在;以激光共聚焦荧光显微镜结合荧光标记物示踪检测,发现不透膜荧光物质分子量为457Da 的萤黄(Lucifer yellow,LYCH),可以共质体运输方式进入存活细胞内,论证了类外连丝这一胞间连丝特定修饰态的存在,并可在一段时间内继续保持生理活性,起到进行物质共质运输的功能。     

蒜鳞片休眠进程中胞间联络的变化及类外连丝结构与功能的研究

利用电子显微镜术,对蒜休眠进程中鳞片薄壁细胞间胞间联络的特征进行了实验观察,发现不同时期胞间联络具有随细胞间生理关系密切程度而呈现相应结构变化的特点。并观察到萌芽期鳞片中衰败细胞与存活细胞之间有类外连丝型胞间连丝的存在;以激光共聚焦荧光显微镜结合荧光标记物示踪检测,发现不透膜荧光物质分子量为457Da的萤黄(Lucifer yellow,LYCH),可以共质体运输方式进入存活细胞内,论证了类外连丝这一胞间连丝特定修饰态的存在,并可在一段时间内继续保持生理活性,起到进行物质共质运输的功能。     

Symplastic communication in the root cap directs auxin distribution to modulate root development

Root cap not only protects root meristem, but also detects and transduces the signals of environmental changes to affect root development. The symplastic communication is an important way for plants to transduce signals to coordinate the development and physiology in response to the changing enviroments. However, it is unclear how the symplastic communication between root cap cells affects root growth. Here we exploit an inducible system to specifically block the symplastic communication in the root cap. Transient blockage of plasmodesmata (PD) in differentiated collumella cells severely impairs the root development in Arabidopsis, in particular in the stem cell niche and the proximal meristem. The neighboring stem cell niche is the region that is most sensitive to the disrupted symplastic communication and responds rapidly via the alteration of auxin distribution. In the later stage, the cell division in proximal meristem is inhibited, presumably due to the reduced auxin level in the root cap. Our results reveal the essential role of the differentiated collumella cells in the root cap mediated signaling system that directs root development.     

Aluminum-induced 1-->3-beta-D-glucan inhibits cell-to-cell trafficking of molecules through plasmodesmata. A new mechanism of aluminum toxicity in plants

Symplastic intercellular transport in plants is achieved by plasmodesmata (PD). These cytoplasmic channels are well known to interconnect plant cells to facilitate intercellular movement of water, nutrients, and signaling molecules including hormones. However, it is not known whether Al may affect this cell-to-cell transport process, which is a critical feature for roots as organs of nutrient/water uptake. We have microinjected the dye lucifer yellow carbohydrazide into peripheral root cells of an Al-sensitive wheat (Triticum aestivum cv Scout 66) either before or after Al treatment and followed the cell-to-cell dye-coupling through PD. Here we show that the Al-induced root growth inhibition is closely associated with the Al-induced blockage of cell-to-cell dye coupling. Immunofluorescence combined with immuno-electron microscopic techniques using monoclonal antibodies against 1-->3-beta-D-glucan (callose) revealed circumstantial evidence that Al-induced callose deposition at PD may responsible for this blockage of symplastic transport. Use of 2-deoxy-D-glucose, a callose synthesis inhibitor, allowed us to demonstrate that a reduction in callose particles correlated well with the improved dye-coupling and reduced root growth inhibition. While assessing the tissue specificity of this Al effect, comparable responses were obtained from the dye-coupling pattern in tobacco (Nicotiana tabacum) mesophyll cells. Analyses of the Al-induced expression of PD-associated proteins, such as calreticulin and unconventional myosin VIII, showed enhanced fluorescence and co-localizations with callose deposits. These results suggest that Al-signal mediated localized alterations to calcium homeostasis may drive callose formation and PD closure. Our data demonstrate that extracellular Al-induced callose deposition at PD could effectively block symplastic transport and communication in higher plants.     

Real-time imaging of phloem unloading in the root tip of Arabidopsis

Confocal laser scanning microscopy (CLSM) has been used to image phloem transport and unloading in the root tip of Arabidopsis. The fluorescent probe 5(6) carboxyfluorescein (CF) was ester loaded into a single cotyledon and the entire seedling placed within an observation chamber under the microscope. Translocation of CF to the root tip was rapid, followed by unloading into discrete concentric files of cells. The position of the prominent unloading ‘zone’ corresponded precisely with that of the two protophloem files of sieve elements, demonstrating a functional role of these cells in symplastic sieve-element unloading. Symplastic transport following unloading was confined to the elongating zone of the root with little basipetal transport to more mature cells. Following photobleaching of the unloading zone, phloem transport was restored immediately into the protophloem sieve elements, followed rapidly by lateral, symplastic sieve-element unloading. The results demonstrate that phloem transport processes can now be imaged in real time, and non-invasively, within an intact plant system.     

Lead accumulation and its translocation barriers in roots of Allium cepa L.—autoradiographic and ultrastructural studies

Abstract Lead migrating through the tissues of Allium cepa L. was found, by electron microscopy, autoradiography and other methods, to encounter at least three barriers to penetration. The layers of protoderm and hypodermic meristematic cells in the root meristematic zone and the layer of endodermis in the mature root zone were barriers to apoplastic transport. The central zone was a barrier to apoplastic and symplastic transport. It comprises the quiescent centre in the root meristem and the central part of the root cap. The cells of the deepest ground meristematic tissue layers seemed to act as a barrier, which keeps lead away from the procambium. Lead accumulated in roots but it was not uniformly distributed between their various tissues. The largest amount of lead accumulated both in ground meristematic and cortex tissues.     

白叶枯病菌毒素对水稻根冠细胞微丝分布的影响

用白叶枯病茵株Ah28产生的毒素,处理水稻野败型细胞质雄性不育系珍汕97A,以异硫氰酸荧光素一鬼笔环肽为探针,观察了毒素处理前后微丝骨架的形态及分布变化。结果显示,毒素处理后,珍汕97A的根冠细胞微丝的密集分布被破坏,纤丝状微丝不复存在,微丝集聚成粗束,有的呈碎片状。说明毒素影响微丝的分布,并探讨了毒素致病机理及微丝参与信号转导的可能性。     

玉米根冠脱落细胞中微丝分布的荧光显微观察(简报)

在对玉米等主要农作物进行病理及致病毒素毒理分析时,经常选用根冠脱落细胞作为试材。许多学眷认为这些脱落的细胞处于垂死状态,然而,后来的一些研究肯定了Knudson的脱落根冠细胞具有活力的观点。Copoorali无菌培养玉米根冠脱落细胞,发现它们可以分裂;Vermeer等提出存活于玉米根际周围的根冠脱落细胞是根系统范围的扩展,它们在土壤中的作用应予以特别注意;Hawes不仅认为它们为活细胞,而且以它们做实验材料用于毒理分析;Guinal对玉米根冠脱落细胞的来源、细胞结构及生理特性进行了详细研究,肯定了细胞     

A MORPHOMETRIC ANALYSIS OF CELLULAR DIFFERENTIATION IN THE ROOT CAP OF ZEA MAYS

In order to quantify the ultrastructural changes associated with cellular differentiation, we have performed a morphometric analysis of the ultrastructure of the calyptrogen, columella, and peripheral cells of the root cap of Zea mays. The relative volumes of the nucleus, nucleolus, and mitochondria in the protoplasm gradually decrease as a cell moves through the root cap. The relative volume of plastids increases 240% during the differentiation of calyptrogen cells into columella cells. This increase is transient, however, since the relative volume of plastids as well as starch in plastids decreases markedly as columella cells differentiate into peripheral cells. Dictyosomes and spherosomes increase more gradually than plastids, peaking in relative volume in the innermost peripheral cells (PCI). The relative volume of the vacuome decreases as calyptrogen cells differentiate into columella cells, after which it increases during the differentiation of peripheral cells. By the time the outermost peripheral cells (PCIII) are sloughed from the cap, the relative volume of the vacuome has almost tripled. These results indicate that each cell type comprising the root cap of Zea mays is characterized by a distinctive ultrastructure. Furthermore, the ultrastructural changes associated with the differentiation of these cells are organelle specific. The results of this study are discussed relative to the function of the various cell types of the root cap.