植物根系质外体屏障研究进展

根是植物吸收水分和矿质营养以维持生命活动的重要器官。根系的构型和超微结构具有物种特异性, 对水分和矿质营养的吸收有不同程度的影响。其中, 内、外皮层的木栓层和凯氏带是2种重要的质外体屏障, 可非定向地阻断水分和离子运输, 在植物生长发育及响应逆境胁迫中发挥重要作用。尽管如此, 植物根系质外体屏障的结构、化学组成、生理功能、生物合成及其调控仅在模式植物拟南芥(Arabidopsis thaliana)中被广泛研究。近年来, 关于作物大麦(Hordeum vulgare)、水稻(Oryza sativa)以及部分牧草的根系质外体屏障研究报道逐渐增多。该文系统比较了拟南芥、大麦、水稻以及部分牧草根系质外体屏障的异同, 提出今后的研究方向, 以期为深入探索禾本科作物和牧草根系质外体屏障在生长发育和逆境适应中的作用奠定理论基础, 并为作物和牧草育种工作提供新思路。     

双穗雀稗(Paspalum distichum)通透性生理和茎解剖结构补充研究

双穗雀稗根外皮层,茎角质层和茎节中的质外体屏障结构阻挡黄连素示踪液透过植物体。茎中机械组织包括周缘厚壁机械组织层,厚壁组织层和维管系统,髓部和皮层的蜂窝状厚角组织。茎中通气组织包括茎节间髓部和皮层的蜂窝状通气组织,茎节内的通气组织。双穗雀稗茎节间具有外侧、内侧和维管系统的质外体屏障,以及茎节周围质外体屏障的封闭结构。因此,该植物体完善的机械组织、通气组织、质外体屏障结构及其离子不通透性是其适应湿地环境的重要结构。     

入侵植物水花生解剖学和组织化学染色研究

水花生(Alternanthera philoxeroides)因其表型可塑性、高生长速率和快速无性繁殖能适应水、陆生境。该文利用光学显微镜和荧光显微镜对水、陆生境的水花生不定根、茎解剖结构、组织化学特征及质外体通透性进行了研究。结果表明:(1)水生境下,其不定根皮层中具较大裂生型通气组织,无次生生长,内皮层具凯氏带且栓质化,皮层和皮下层明显木质化。(2)在陆生环境下,其不定根有次生生长,胞间具通气组织,内皮层具凯氏带且栓质化,皮层和皮下层略木质化;此外,不定根还具额外形成层,产生次生维管束、薄壁组织和不定芽;多年生不定根中具直接分裂的薄壁组织,周皮具凯氏带,且栓质化和木质化。(3)水、陆生境下,其匍匐茎具髓和中空髓腔,发生次生生长,具裂-溶生型通气组织、单层内皮层、厚角组织和木质化且栓质化的角质层,陆生匍匐茎周皮栓质化且木质化。(4)水花生质外体屏障结构组成复杂,黄连素无法穿透质外体屏障结构。水花生的上述解剖学特征,是水花生适应水、陆生境的有力证据。     

菰适应湿地环境的解剖和屏障结构特征研究

菰(Zizania latifolia)是一种多年生挺水植物,为了探讨该植物根、茎和叶的解剖结构、组织化学及其质外体屏障的通透性生理。该文利用光学显微镜和荧光显微镜,对菰的根、茎、叶进行了解剖学和组织化学研究。结果表明:(1)菰不定根解剖结构由外而内分别为表皮、外皮层、单层细胞的厚壁机械组织层、皮层、内皮层和维管柱;茎结构由外而内分别为角质层、表皮、周缘厚壁机械组织层、皮层、具维管束的厚壁组织层和髓腔。叶鞘具有表皮和具维管束皮层,叶片具有表皮,叶肉和维管束。(2)不定根具有位于内侧的内皮层及其邻近栓质化细胞和外侧的外皮层组成的屏障结构;茎具内侧厚壁机械组织层,外侧的角质层和周缘厚壁机械组织层组成的屏障结构,屏障结构的细胞壁具凯氏带、木栓质和木质素沉积的组织化学特点,叶表面具有角质层。(3)菰通气组织包括根中通气组织,茎、叶皮层的通气组织和髓腔。(4)菰的屏障结构和解剖结构是其适应湿地环境的重要特征,但其茎周缘厚壁层和厚壁组织层较薄。由此推测,菰适应湿地环境,但在旱生环境中分布有一定的局限性。     

植物凯氏带形成分子机制及功能特点的研究进展

凯氏带位于被子植物初生根内皮层细胞,环绕细胞1周,是与质膜紧密结合的非极性带状增厚结构。凯氏带作为植物根中离子径向运输障碍,调节离子的质外体吸收途径,迫使土壤中的离子通过内皮层细胞膜,选择性地进入中柱。凯氏带发现于1865年,但直至拟南芥凯氏带蛋白的发现和凯氏带阻滞作用物质基础被揭示,凯氏带的形成机理和功能才逐渐为人们所认知。凯氏带的物质基础为木质素,其形成需要由凯氏带蛋白和受体激酶组成的合成平台。细胞内部的木质素单体经ABCG载体运输到凯氏带的形成区,经ESB1dirigent蛋白、RBOHF氧化酶和PER64过氧化物酶等催化,合成木质素。该文对近年来国内外有关凯氏带形成的分子机制和功能特点方面的研究进展进行综述,为进一步理解和解析凯氏带的形成机理和功能提供参考。     

单子叶植物根的内皮层及其观察方法

裸子植物和具有少量次生长的双子叶植物根的内皮层细胞通常都具有径向壁和横壁加厚而成的凯氏带结构。但多数单子叶植物和少数双子叶植物(没有次生生长的种类)根的内皮层细胞只有在发育的早期(幼根)具有凯氏带,发育的后期(较老的根)内皮层细胞壁内敷贴了一整层栓质层,随后细胞壁又经过了五面加厚(只有邻接皮层一面的壁没有加厚)或六面加厚(即全部细胞壁都加厚),加厚部分都经过了木质化,因此这时就看不到凯氏带了。由于内皮层细胞内敷贴了一整层栓质层,这类植物根的维管柱似乎被一层不透水的套子所隔开了。实际不然,因为并不是全部内皮层细胞都具有五面或六面加厚的壁,通常有一部分内皮层细胞,就是在根的横切面上靠近木质部角端的那一部分内皮层细胞,仍有未栓质化、未加厚、只有凯氏带的纤维素薄壁,这一部分内皮层细胞     

植物凯氏带化学成分、生理功能及相关调控机制的研究进展

凯氏带是一种环绕在内皮层细胞径向壁和横向壁上的木质化和木栓化带状增厚结构.凯氏带作为一种保护性屏障,不仅可以调节物质和水分在维管束与皮层细胞之间的渗透性流动,而且在抵御盐胁迫、生物胁迫以及营养物质的选择性吸收等方面也发挥重要作用.随着近年来生物化学分析技术及遗传学研究方法的发展,凯氏带的相关研究取得了极大进展.本文简要概述了凯氏带在植物体内的组织结构、化学组成和发育过程,并重点对凯氏带生理功能、重要调控基因及其相关调控机制等方面的最新进展进行了总结,以期为凯氏带的深入研究和开发利用提供参考.     

植物凯氏带的研究进展

凯氏带是植物内皮层细胞径向壁和横向壁的带状增厚部分 ,近年来对其细微结构、化学成分、生理功能以及在植物体的存在部位等方面的研究工作已成为国内、外的研究热点。本文就凯氏带上述诸方面的研究进展 ,作一详尽的论述 ,同时还着重对凯氏带研究的新方法和新技术作了评述。最后对凯氏带的进一步研究工作提供新的启示     

重金属镉(Cd)在植物体内的转运途径及其调控机制

重金属镉(Cd)的毒害效应与其由土壤向植物地上部分运输有关,揭示Cd~(2+)转运途径及其调控机制可为提高植物抗镉性以及镉污染的植物修复提供依据。对Cd~(2+)在植物体内的转运途径,特别是限制Cd~(2+)移动的细胞结构和分子调控机制研究进展进行了回顾。Cd~(2+)通过共质体和质外体途径穿过根部皮层进入木质部的过程中,大部分在皮层细胞间沉积,少部分抵达中柱后转移到地上部分。为了免受Cd~(2+)的危害,植物体产生了多种限制Cd~(2+)吸收和转移的生理生化机制:1)环绕在内皮层径向壁和横向壁上的凯氏带阻止Cd~(2+)以质外体途径进入木质部;2)螯合剂与进入根的Cd~(2+)螯合形成稳定化合物并区隔在液泡中;3)通过H+/Cd~(2+)离子通道等将Cd~(2+)逆向转运出根部。植物共质体和质外体途径转运重金属镉的能力以及两条途径的串扰尚待进一步明晰和阐明。     

植物内皮层的分化及其屏障功能研究进展

植物根系最主要的作用之一是从土壤中获取养分并将其运输至地上部。水和营养物质径向穿过根的表皮、皮层、内皮层等所有外部细胞层,才能到达中柱,以供地上部代谢所需。其中,内皮层细胞在发育过程中会经历两个特殊的分化阶段,分别形成凯氏带和木栓层两种扩散屏障,二者在控制养分获取与流失方面起着重要的作用。该文就近年来国内外有关植物内皮层分化过程及其屏障功能方面的研究进展进行了综述,以期对深入探索内皮层屏障在植物生长发育和逆境适应中的作用提供参考,为植物育种工作开辟新的思路。     

The exodermis: a variable apoplastic barrier.

The exodermis (hypodermis with Casparian bands) of plant roots represents a barrier of variable resistance to the radial flow of both water and solutes and may contribute substantially to the overall resistance. The variability is a result largely of changes in structure and anatomy of developing roots. The extent and rate at which apoplastic exodermal barriers (Casparian bands and suberin lamellae) are laid down in radial transverse and tangential walls depends on the response to conditions in a given habitat such as drought, anoxia, salinity, heavy metal or nutrient stresses. As Casparian bands and suberin lamellae form in the exodermis, the permeability to water and solutes is differentially reduced. Apoplastic barriers do not function in an all-or-none fashion. Rather, they exhibit a selectivity pattern which is useful for the plant and provides an adaptive mechanism under given circumstances. This is demonstrated for the apoplastic passage of water which appears to have an unusually high mobility, ions, the apoplastic tracer PTS, and the stress hormone ABA. Results of permeation properties of apoplastic barriers are related to their chemical composition. Depending on the growth regime (e.g. stresses applied) barriers contain aliphatic and aromatic suberin and lignin in different amounts and proportion. It is concluded that, by regulating the extent of apoplastic barriers and their chemical composition, plants can effectively regulate the uptake or loss of water and solutes. Compared with the uptake by root membranes (symplastic and transcellular pathways), which is under metabolic control, this appears to be an additional or compensatory strategy of plants to acquire water and solutes.     

Casparian strip development and its potential function in salt tolerance

The root system is particularly affected by unfavourable conditions because it is in direct contact with the soil environment. Casparian strips, a specialised structure deposited in anticlinal walls, are characterised by the impregnation of the primary wall pores with lignin and suberin. The Casparian strips in the endo- and exodermis of vascular plant roots appear to play an important role in preventing the non-selective apoplastic bypass of salts into the stele along the apoplast under salt stress. However, only a few investigations have examined the deposition and function of these apoplastic barriers in response to salt stress in higher plants.     

Net sodium fluxes change significantly at anatomically distinct root zones of rice (Oryza sativa L.) seedlings

Casparian bands of endodermis and exodermis play crucial roles in blocking apoplastic movement of ions and water into the stele of roots through the cortex. These apoplastic barriers differ considerably in structure and function along the developing root. The present study assessed net Na⁺ fluxes in anatomically distinct root zones of rice seedlings and analyzed parts of individual roots showing different Na⁺ uptake. The results indicated that anatomically distinct root zones contributed differently to the overall uptake of Na⁺. The average Na⁺ uptake in root zones in which Casparian bands of the endo- and exo-dermis were interrupted by initiating lateral root primordia (root zone III) was significantly greater than that at the root apex, where Casparian bands were not yet formed (root zone I), or in the region where endo- and exo-dermis with Casparian bands were well developed (root zone II). The measurement of net Na⁺ fluxes using a non-invasive scanning ion-selective electrode technique (SIET) demonstrated that net Na⁺ flux varied significantly in different positions along developing rice roots, and a net Na⁺ influx was obvious at the base of young lateral root primordia. Since sodium fluxes changed significantly along developing roots of rice seedlings, we suggest that the significantly distinct net Na⁺ flux profile may be attributed to different apoplastic permeability due to lateral root primordia development for non-selective apoplastic bypass of ions along the apoplast.     

白皮松针叶内皮层在盐胁迫中的屏障作用研究

应用荧光显微技术、傅里叶变换显微红外光谱分析(FTIR)、扫描电镜及X-射线能谱微区分析等手段,对白皮松(Pinus bungeana)子叶、初生叶及2a生针叶内皮层细胞径向壁的显微结构特征、化学成分,以及在叶子横切面上Na和Cl的微区分布进行分析。通过荧光显微观察发现,白皮松子叶内皮层不具凯氏带,而初生叶及2a生针叶均存在凯氏带加厚现象。根据FTIR的检测结果显示:子叶内皮层细胞径向壁不含木栓质或极少,2a生针叶内皮层细胞径向壁木栓质含量高于初生叶。对相应区域的X射线微区分析表明,子叶内皮层对Na和Cl在质外体运输中不起障碍作用,而初生叶与2a生针叶内皮层阻碍Na和Cl以质外体途径进入维管组织。研究结果表明:具凯氏带加厚的内皮层细胞壁中木栓质含量决定其在质外体运输过程中的生理功能。     

Casparian strips in needles are more solute permeable than endodermal transport barriers in roots of Pinus bungeana

The structure and development of endodermal Casparian strips in Pinus bungeana needles and roots were studied by scanning electron microscopy and fluorescence microscopy. Primary pit fields (PFs) frequently occurred in radial walls of Casparian strips isolated from needles, whereas PFs were never detected in Casparian strips from roots. Formation of Casparian strips in needles as well as roots started at the outer parts of the radial walls and they finally occupied the entire radial walls of the endodermis. Fourier transform infrared (FTIR) spectroscopy of Casparian strips isolated from roots revealed significant absorption bands characteristic for suberin. However, in Casparian strips of needles, evidence for suberin was rarely detected by FTIR spectroscopy. The apoplastic permeability of Casparian strips in needles and roots was probed by the apoplastic tracers calcofluor and berberine. Casparian strips in roots efficiently blocked the apoplastic transport (AT) of calcofluor and berberine. Casparian strips in needles blocked the AT of calcofluor, but diffusion of berberine was not inhibited and berberine thiocyanate crystals were detectable in the vascular tissue of the needles. From the data presented, it must be concluded that Casparian strips in needles, which are characterized by the absence of suberin, are more solute permeable compared with Casparian strips in roots.     

黄连体内黄连素的组织器官定位和根尖屏障结构特征研究

黄连(Coptis chinensis)是毛茛科著名药材,该文研究了黄连体内黄连素在组织器官中的分布规律和根尖屏障结构特征。在白光和荧光显微镜下,组织器官中黄连素在蓝色激发光下自发黄色荧光,黄连素-苯胺兰对染研究细胞壁凯氏带和木质化,苏丹7B染色栓质层,间苯三酚-盐酸染色木质化。结果表明:黄连不定根初生结构为维管柱、内皮层、皮层、外皮层和表皮组成;次生结构以次生木质部为主、次生韧皮部和木栓层组成。黄连根茎初生结构由角质层,皮层和维管柱组成;次生结构由木栓层、皮层和维管柱组成,以皮层和维管柱为主。叶柄结构为髓、含维管束的厚壁组织层、皮层和角质层。黄连不定根的屏障结构初生结构时期由栓质化和木质化的内皮层、外皮层;次生结构时期为木栓层组成;根状茎的为角质层和木栓层。黄连素主要沉积分布在不定根和茎的木质部,叶柄的厚壁组织层,木质部和厚壁组织是鉴别黄连品质的重要部位。黄连根尖外皮层及早发育,同时初生木质部有黄连素沉积结合,可能造成水和矿质吸收和运输的阻碍,也是黄连适应阴生环境的重要原因。     

Analytical electron microscopical investigations on the apoplastic pathways of lanthanum transport in barley roots

 In transmission electron microscopy studies, lanthanum ions have been used as electron-opaque tracers to delineate the apoplastic pathways for ion transport in barley (Hordeum vulgare L.) roots. To localize La³⁺ on the subcellular level, e.g. in cell walls and on the surface of membranes, electron-energy-loss spectroscopy and electron-spectroscopic imaging were used. Seminal and nodal roots were exposed for 30 min to 1 mM LaCl₃ and 10 mM LaCl₃, respectively. In seminal roots, possessing no exodermis, La³⁺ diffusion through the apoplast was stopped by the Casparian bands of the endodermis. In nodal roots with an exodermis, however, La³⁺ diffusion through the cortical apoplast had already stopped at the tight junctions of the exodermal cell walls resembling the Casparian bands of the endodermis. Therefore, we conclude that in some specialized roots such as the nodal roots of barley, the physiological role of the endodermis is largely performed by the exodermis. Received: 28 July 1999 / Accepted: 24 February 2000     

Evidence for symplastic involvement in the radial movement of calcium in onion roots

The pathway of Ca(2+) movement from the soil solution into the stele of the root is not known with certainty despite a considerable body of literature on the subject. Does this ion cross an intact, mature exodermis and endodermis? If so, is its movement through these layers primarily apoplastic or symplastic? These questions were addressed using onion (Allium cepa) adventitious roots lacking laterals. Radioactive Ca(2+) applied to the root tip was not transported to the remainder of the plant, indicating that this ion cannot be supplied to the shoot through this region where the exodermis and endodermis are immature. A more mature zone, in which the endodermal Casparian band was present, delivered 2.67 nmol of Ca(2+) mm(-1) treated root length d(-1) to the transpiration stream, demonstrating that the ion had moved through an intact endodermis. Farther from the root tip, a third zone in which Casparian bands were present in the exodermis as well as the endodermis delivered 0.87 nmol Ca(2+) mm(-1) root length d(-1) to the transpiration stream, proving that the ion had moved through an unbroken exodermis. Compartmental elution analyses indicated that Ca(2+) had not diffused through the Casparian bands of the exodermis, and inhibitor studies using La(3+) and vanadate (VO(4)(3-)) pointed to a major involvement of the symplast in the radial transport of Ca(2+) through the endodermis. It was concluded that in onion roots, the radial movement of Ca(2+) through the exodermis and endodermis is primarily symplastic.     

Abscisic acid‐mediated modifications of radial apoplastic transport pathway play a key role in cadmium uptake in hyperaccumulator Sedum alfredii

Abscisic acid (ABA) is a key phytohormone underlying plant resistance to toxic metals. However, regulatory effects of ABA on apoplastic transport in roots and consequences for uptake of metal ions are poorly understood. Here, we demonstrate how ABA regulates development of apoplastic barriers in roots of two ecotypes of Sedum alfredii and assess effects on cadmium (Cd) uptake. Under Cd treatment, increased endogenous ABA level was detected in roots of nonhyperaccumulating ecotype (NHE) due to up‐regulated expressions of ABA biosynthesis genes (SaABA2, SaNCED), but no change was observed in hyperaccumulating ecotype (HE). Simultaneously, endodermal Casparian strips (CSs) and suberin lamellae (SL) were deposited closer to root tips of NHE compared with HE. Interestingly, the vessel‐to‐CSs overlap was identified as an ABA‐driven anatomical trait. Results of correlation analyses and exogenous applications of ABA/Abamine indicate that ABA regulates development of both types of apoplastic barriers through promoting activities of phenylalanine ammonialyase, peroxidase, and expressions of suberin‐related genes (SaCYP86A1, SaGPAT5, and SaKCS20). Using scanning ion‐selected electrode technique and PTS tracer confirmed that ABA‐promoted deposition of CSs and SL significantly reduced Cd entrance into root stele. Therefore, maintenance of low ABA levels in HE minimized deposition of apoplastic barriers and allowed maximization of Cd uptake via apoplastic pathway.