植物角质膜研究进展

植物的角质膜是植物与外界环境的交界面, 有利于植物减少蒸腾、抵抗紫外伤害和防止病虫害等。植物与外部环境相互作用的过程中, 其角质膜会构建自身的防御系统, 如通过自身结构、成分的改变及产生次生代谢产物来减轻外界不利环境因子的胁迫。因此, 植物角质膜与环境之间的密切关系对于植物生长有着重要意义。该文综述了植物角质膜的结构、成分、形成、功能及其与次生代谢和环境的关系, 同时对研究中存在的问题进行了讨论并展望了相关领域的研究前景。     

植物角质膜的结构、组成和生物学功能研究进展

植物角质膜是覆盖在植物最外层的一类有机混合物的总称,它是植物抵抗外界环境刺激的最后一道屏障,在植物生长发育过程中起重要作用。该文总结了近几十年来国内外关于角质膜的研究进展,通过植物角质膜的形态结构、化学组成、生物学功能等几个方面对角质膜的研究状况做系统综述,探讨目前研究中存在的一些问题,展望角质膜研究前景,以期为从事角质膜相关领域的研究提供可借鉴的参考依据。     

3种芦荟属植物叶的表皮扫描电镜观察及芦荟素含量的测定

以库拉索芦荟、木立芦荟和皂质芦荟为材料,用扫描电镜观察其叶表皮气孔和角质膜的结构,用高效液相色谱法(HPLC)测定了3种芦荟属植物叶中芦荟素的含量。扫描电镜观察结果表明,3种芦荟叶表皮都覆盖有厚的角质膜,气孔下陷,表现出典型的旱生植物特征。但角质膜的纹饰和厚度在不同芦荟间有显着差异。木立芦荟角质膜表面呈瘤状突起,角质膜厚度为5～6μm,库拉索芦荟和皂质芦荟的角质膜表面较平,库拉索芦荟的角质膜厚度为3～4μm,皂质芦荟的角质膜厚度为8～10μm。高效液相色谱法(HPLC)测定结果表明,木立芦荟叶含芦荟素最高,库拉索芦荟叶含量较低,而皂质芦荟叶含量最低。此外,本文还初步探讨了芦荟属植物叶表皮结构与芦荟素含量的关系。     

中国桑寄生科植物叶表皮微形态

通过扫描电镜对中国桑寄生科桑寄生亚科8属18种和槲寄生亚科1属2种植物成熟叶的上、下叶表皮内表面和下表皮外表面进行了研究。内面观发现桑寄生科植物叶上、下表皮形状为多边形,垂周壁式样平直或稍弓形,常具有角质增厚,平周壁常覆盖厚角质或颗粒状、丝状角质增厚;气孔存在于上下表皮,通常下表皮较多,气孔的形状,特别是保卫细胞的形态在亚科间、属间或种间都具有一定的差异,气孔器类型为平列型或单圈型。下表皮表面观察了的角质膜和蜡质纹饰、气孔的形状,外部气孔缘及外部气孔缘内缘的特征。这些特征在亚科或属级水平上较为稳定,有的也表现出种间差异,有一定的分类价值。从气孔形态和外部气孔周围角质膜来看,两亚科显示出明显的不同:桑寄生亚科上、下表皮均具有内部气孔缘,而槲寄生亚科没有此结构;桑寄生亚科外部气孔周围角质膜增厚成环状,其上具增厚的条纹,而槲寄生亚科外部气孔周围角质膜增厚成脊状,不具条纹。这些特征支持槲寄生亚科作为独立1个科来处理。     

植物角质蒸腾的几个方面

角质蒸腾速率和植物生态型有密切关系,与普通品种相比抗旱的高粱和玉米品种角质蒸腾较弱,或差异不明显,但节水途径不同。强光促进角质层腊质的形成,使角质蒸腾减弱。高湿度下生长的植物移到湿度较低处后,其角质蒸腾与总蒸腾速率都明显增大。土壤干旱对角质蒸腾的影响因植物种类和水分胁迫的具体情况而异。角质层腊质含量和角质蒸腾速率一般呈反相关。     

安徽景天属（Sedum）植物叶表皮研究

利用光学显微镜和扫描电子显微镜观察安徽省１２种景天属（Ｓｅｄｕｍ）植物的叶表皮,统计并测量报气孔类型,气孔大小,气孔密度及气孔指数等,描述了表皮角质膜,蜡质纹饰及气孔外拱盖的有关特征。结果表明：景天属植物叶表皮气孔器为不等细胞型。气孔器类型,表皮细胞形状及垂周壁式样,角质膜,蜡质纹饰等,种间无差异或极小,而气孔大小,气孔器分布特征,气孔密度,表皮毛等种间差异较大,建议在种间区分上将差异大的特征作为     

五种海桑属红树植物叶片的结构及其生态适应

对海桑属五种红树植物的叶片进行解剖学的观察和研究。结果表明:海桑属红树植物的叶片上、下均有栅栏组织,栅栏组织与叶片厚的比值在0.3375～0.4349之间,属等面叶;表皮的角质膜厚1.88～7.63μm;输导组织发达;多数种类的栅栏组织中有分泌腔,而海绵组织中有单宁异细胞分布。说明了海桑属红树植物叶片的结构具有很强的耐旱及抗腐蚀能力。     

植物寄生线虫侧带的制作方法

植物寄生线虫体形较小,长度一般不超过1—2毫米,宽度约为0.03—0.05毫米。线虫体外皆具有一层非细胞结构的弹性角质膜,其上常有纵纹和环纹,以及其他形状的花纹。在线虫两侧常有数条由虫体前端直达后端的纵纹组成的侧带(Lateral field)。这些角质膜上的花纹     

植物叶表面的附着物

植物叶表面上的附着物实际上是植物叶表面的分泌物。很多物质可以从原生质体中分离并沉积下来,这实际上是一种分泌过程,这些沉积物质就是分泌物。叶表皮细胞外壁上附着的角质膜、蜡质以及各种盐的     

鹿药属植物叶表皮特征及其系统学意义

采用光学显微镜和扫描电镜对鹿药属12种植物的叶表皮进行了观察,首次报道了12种鹿药属(Sm ilaci-na)植物叶表皮的微形态特征。结果表明:气孔器普遍存在于叶的下表皮,少数种的上表皮也有分布,均为不规则形。叶表皮细胞形状为多边形或不规则形,垂周壁式样可区分为近平直、浅波状和波状。在扫描电镜下,叶表皮气孔器外拱盖内缘为近平滑、浅波状或波状;角质膜条纹状,有的条纹隆起,有的条纹上附有颗粒和晶簇。气孔器的分布、气孔器外拱盖内缘形态以及角质膜等特征对该属部分种的区分具有一定的参考价值。     

植物角质层蜡质基因的研究进展

角质层是覆盖在植物地上部分最表层的保护层,具有降低植物表面的水分散失、防止紫外线辐射伤害和抵抗病虫害侵入等环境胁迫等功能,在植物适应外界环境作用方面起重要作用.作对近年来角质层蜡质基因的研究进展进行综述,同时也对蜡质基因的研究前景提出一些看法.     

Disruptions of the Arabidopsis Enoyl-CoA reductase gene reveal an essential role for very-long-chain fatty acid synthesis in cell expansion during plant morphogenesis

In the absence of cell migration, plant architecture is largely determined by the direction and extent of cell expansion during development. In this report, we show that very-long-chain fatty acid (VLCFA) synthesis plays an essential role in cell expansion. The Arabidopsis thaliana eceriferum10 (cer10) mutants exhibit severe morphological abnormalities and reduced size of aerial organs. These mutants are disrupted in the At3g55360 gene, previously identified as a gene coding for enoyl-CoA reductase (ECR), an enzyme required for VLCFA synthesis. The absence of ECR activity results in a reduction of cuticular wax load and affects VLCFA composition of seed triacylglycerols and sphingolipids, demonstrating in planta that ECR is involved in all VLCFA elongation reactions in Arabidopsis. Epidermal and seed-specific silencing of ECR activity resulted in a reduction of cuticular wax load and the VLCFA content of seed triacylglycerols, respectively, with no effects on plant morphogenesis, suggesting that the developmental phenotypes arise from abnormal sphingolipid composition. Cellular analysis revealed aberrant endocytic membrane traffic and defective cell expansion underlying the morphological defects of cer10 mutants.     

ABC-type transporters and cuticle assembly: Linking function to polarity in epidermis cells

The aerial organs of plants are covered with a cuticle, a continuous layer overlaying the outermost cell walls of the epidermis. The cuticle is composed of two major classes of the lipid biopolymers: cutin and waxes, collectively termed cuticular lipids. Biosynthesis and transport of cuticular lipids occur predominantly in the epidermis cells. In the transport pathway, cuticular lipids are exported from their site of biosynthesis in the ER/plastid to the extracellular space through the plasma membrane and cell wall. Growing evidence suggests that ATP-binding cassette (ABC) transporters are implicated in transport of cuticular lipids across the plasma membrane of epidermal cells. The Arabidopsis ABC-type transporter protein CER5 (WBC12) was reported to act as a wax monomers transporter. In recent works, our group and others showed that a CER5-related protein, DESPERADO (DSO/WBC11), is required for cutin and wax monomers transport through the plasma membrane of Arabidopsis epidermis cells. Unlike the cer5 mutant, DSO loss-of-function had a profound effect on plant growth and development, particularly dwarfism, postgenital organ fusions, and altered epidermal cell differentiation. The partially overlapping function of CER5 and DSO and the fact that these proteins are half-size ABC transporters suggest that they might form a hetero-dimeric complex while transporting wax components. An intriguing observation was the polar localization of DSO in the distal part of epidermis cells. This polar expression might be explained by DSO localization within lipid rafts, specific plasma membrane microdomains which are associated with polar protein expression. In this review we suggest possible mechanisms for cuticular lipids transport and a link between DSO function and polar expression. Furthermore, we also discuss the subsequent transport of cuticular constituents through the hydrophobic cell wall and the possible involvement of lipid transfer proteins in this process.Key words: ABC transporter, cuticular lipids, polar expression, plasma membrane, epidermis     

Host surface properties affect prepenetration processes in the barley powdery mildew fungus

The initial contact between Blumeria graminis f.sp. hordei and its host barley (Hordeum vulgare) takes place on epicuticular waxes at the surfaces of aerial plant organs. Here, the extent to which chemical composition, crystal structure and hydrophobicity of cuticular waxes affect fungal prepenetration processes was explored. The leaf surface properties of barley eceriferum (cer) wax mutants were characterized in detail. Barley leaves and artificial surfaces were used to investigate the early events of fungal infection. Even after epicuticular waxes had been stripped away, cer mutant leaf surfaces did not affect fungal prepenetration properties. Removal of total leaf cuticular waxes, however, resulted in a 20% reduction in conidial germination and differentiation. Two major components of barley leaf wax, hexacosanol and hexacosanal, differed considerably in their ability to effectively trigger conidial differentiation on glass surfaces. While hexacosanol, attaining a maximum hydrophobicity with contact angles of no more than 80 degrees, proved to be noninductive, hexacosanal significantly stimulated differentiation in c. 50% of B. graminis conidia, but only at contact angles > 80 degrees. These results, together with an observed inductive effect of highly hydrophobic, wax-free artificial surfaces, provide new insights into the interplay of physical and chemical surface cues involved in triggering prepenetration processes in B. graminis.     

Overexpression of Arabidopsis ECERIFERUM1 promotes wax very-long-chain alkane biosynthesis and influences plant response to biotic and abiotic stresses

Land plant aerial organs are covered by a hydrophobic layer called the cuticle that serves as a waterproof barrier protecting plants against desiccation, ultraviolet radiation, and pathogens. Cuticle consists of a cutin matrix as well as cuticular waxes in which very-long-chain (VLC) alkanes are the major components, representing up to 70% of the total wax content in Arabidopsis (Arabidopsis thaliana) leaves. However, despite its major involvement in cuticle formation, the alkane-forming pathway is still largely unknown. To address this deficiency, we report here the characterization of the Arabidopsis ECERIFERUM1 (CER1) gene predicted to encode an enzyme involved in alkane biosynthesis. Analysis of CER1 expression showed that CER1 is specifically expressed in the epidermis of aerial organs and coexpressed with other genes of the alkane-forming pathway. Modification of CER1 expression in transgenic plants specifically affects VLC alkane biosynthesis: waxes of TDNA insertional mutant alleles are devoid of VLC alkanes and derivatives, whereas CER1 overexpression dramatically increases the production of the odd-carbon-numbered alkanes together with a substantial accumulation of iso-branched alkanes. We also showed that CER1 expression is induced by osmotic stresses and regulated by abscisic acid. Furthermore, CER1-overexpressing plants showed reduced cuticle permeability together with reduced soil water deficit susceptibility. However, CER1 overexpression increased susceptibility to bacterial and fungal pathogens. Taken together, these results demonstrate that CER1 controls alkane biosynthesis and is highly linked to responses to biotic and abiotic stresses.     

Estimation of the solubility parameters of model plant surfaces and agrochemicals: a valuable tool for understanding plant surface interactions

Background

Most aerial plant parts are covered with a hydrophobic lipid-rich cuticle, which is the interface between the plant organs and the surrounding environment. Plant surfaces may have a high degree of hydrophobicity because of the combined effects of surface chemistry and roughness. The physical and chemical complexity of the plant cuticle limits the development of models that explain its internal structure and interactions with surface-applied agrochemicals. In this article we introduce a thermodynamic method for estimating the solubilities of model plant surface constituents and relating them to the effects of agrochemicals.

Results

Following the van Krevelen and Hoftyzer method, we calculated the solubility parameters of three model plant species and eight compounds that differ in hydrophobicity and polarity. In addition, intact tissues were examined by scanning electron microscopy and the surface free energy, polarity, solubility parameter and work of adhesion of each were calculated from contact angle measurements of three liquids with different polarities. By comparing the affinities between plant surface constituents and agrochemicals derived from (a) theoretical calculations and (b) contact angle measurements we were able to distinguish the physical effect of surface roughness from the effect of the chemical nature of the epicuticular waxes. A solubility parameter model for plant surfaces is proposed on the basis of an increasing gradient from the cuticular surface towards the underlying cell wall.

Conclusions

The procedure enabled us to predict the interactions among agrochemicals, plant surfaces, and cuticular and cell wall components, and promises to be a useful tool for improving our understanding of biological surface interactions.

     

Monoacylglycerols are components of root waxes and can be produced in the aerial cuticle by ectopic expression of a suberin-associated acyltransferase

The interface between plants and the environment is provided for aerial organs by epicuticular waxes that have been extensively studied. By contrast, little is known about the nature, biosynthesis, and role of waxes at the root-rhizosphere interface. Waxes isolated by rapid immersion of Arabidopsis (Arabidopsis thaliana) roots in organic solvents were rich in saturated C18-C22 alkyl esters of p-hydroxycinnamic acids, but also contained significant amounts of both alpha- and beta-isomers of monoacylglycerols with C22 and C24 saturated acyl groups and the corresponding free fatty acids. Production of these compounds in root waxes was positively correlated to the expression of sn-glycerol-3-P acyltransferase5 (GPAT5), a gene encoding an acyltransferase previously shown to be involved in aliphatic suberin synthesis. This suggests a direct metabolic relationship between suberin and some root waxes. Furthermore, when ectopically expressed in Arabidopsis, GPAT5 produced very-long-chain saturated monoacylglycerols and free fatty acids as novel components of cuticular waxes. The crystal morphology of stem waxes was altered and the load of total stem wax compounds was doubled, although the major components typical of the waxes found on wild-type plants decreased. These results strongly suggest that GPAT5 functions in vivo as an acyltransferase to a glycerol-containing acceptor and has access to the same pool of acyl intermediates and/or may be targeted to the same membrane domain as that of wax synthesis in aerial organs.     

Cloning and characterization of GLOSSY1, a maize gene involved in cuticle membrane and wax production

The cuticle covering the aerial organs of land plants plays a protective role against several biotic and abiotic stresses and, in addition, participates in a variety of plant-insect interactions. Here, we describe the molecular cloning and characterization of the maize (Zea mays) GLOSSY1 (GL1) gene, a component of the pathway leading to cuticular wax biosynthesis in seedling leaves. The genomic and cDNA sequences we isolated differ significantly in length and in most of the coding region from those previously identified. The predicted GL1 protein includes three histidine-rich domains, the landmark of a family of membrane-bound desaturases/hydroxylases, including fatty acid-modifying enzymes. GL1 expression is not restricted to the juvenile developmental stage of the maize plant, pointing to a broader function of the gene product than anticipated on the basis of the mutant phenotype. Indeed, in addition to affecting cuticular wax biosynthesis, gl1 mutations have a pleiotropic effect on epidermis development, altering trichome size and impairing cutin structure. Of the many wax biosynthetic genes identified so far, only a few from Arabidopsis (Arabidopsis thaliana) were found to be essential for normal cutin formation. Among these is WAX2, which shares 62% identity with GL1 at the protein level. In wax2-defective plants, cutin alterations induce postgenital organ fusion. This trait is not displayed by gl1 mutants, suggesting a different role of the maize and Arabidopsis cuticle in plant development.     

Composition of Lipid-derived Polymers from Different Anatomical Regions of Several Plant Species

The composition of the aliphatics of the protective cuticular polymers from different anatomical regions from several plant species was determined by combined gas-liquid chromatography and mass spectrometry of the depolymerization products derived from the polymers. The polymer from the aerial parts of Vicia faba showed similar composition; dihydroxypalmitic acid was the major (>85%) component of the cutin covering leaves, petioles, flower petals and stem with smaller amounts of palmitic acid and ω-hydroxy palmitic acid. On the other hand, the chief components of the polymer from the tap root were ω-hydroxy C_16:0 and C_18:1 acids and/or the corresponding dicarboxylic acids. The positional isomer composition of the dihydroxy C₁₆ acids was shown to be dependent upon anatomical location, developmental stage, and light. Apple cutin from rapidly expanding organs (flower petal and stigma) was shown to contain predominately C₁₆ family acids whereas the C₁₈ family dominated in cutin of slower growing organs (leaf and fruit). The composition of the aliphatic components of cutin found in the seed coats of pea, corn, barley, and lettuce was found to be similar to that of the cuticular polymer of the leaves in each species.     

植物超长链脂肪酸及角质层蜡质生物合成相关酶基因研究现状

超长链脂肪酸(very long chain fatty acids, VLCFAs)在生物体中具有广泛的生理功能, 它们参与种子甘油酯、生物膜膜脂及鞘脂的合成, 并为角质层蜡质的生物合成提供前体物质。角质层是覆盖在植物地上部分最表层的保护层, 由角质和蜡质组成, 其中蜡质又分为角质层表皮蜡和内部蜡, 在植物生长发育、适应外界环境方面起重要作用。VLCFAs的合成由脂肪酰-CoA延长酶催化, 该酶是由b-酮脂酰-CoA合酶、b-酮脂酰-CoA还原酶、b-羟脂酰-CoA脱水酶和反式烯脂酰-CoA还原酶组成的多酶体系。合成后的VLCFAs通过脱羰基与酰基还原作用进入角质层蜡质合成途径, 形成各种蜡质组分。文章就VLCFAs及角质层蜡质合成代谢途径中相关酶基因研究进展方面做了综述, 并对植物蜡质基因研究中存在的问题提出一些看法。