植物表皮蜡质参与干旱胁迫的反应机制

植物表皮是植物与外部环境直接接触的部位,包括具有立体网状结构的角质和填充其间并覆盖其上的蜡质。植物在适应外界环境的过程中,表皮蜡质形成了特殊的结构和复杂的化学组成。植物表皮蜡质最重要的功能是参与阻止植物非气孔性失水,提高植物对水分的利用效率,以实现对干旱环境的适应。干旱环境会导致植物表皮蜡质代谢的变化,这种变化最终通过调控基因表达来实现。目前已经发现了多个蜡质代谢相关基因参与了植物对干旱环境的适应,部分基因已经成功克隆并且用于改良农作物的抗旱性。但这些基因参与干旱响应的分子机制及其与ABA的关系并不很清楚。就植物适应水分胁迫而发生的包括蜡质组成和含量在内的代谢变化,以及该过程中所涉及的主要基因及其分子生物学研究进行综述。探讨表皮蜡质在植物适应干旱中的重要作用及其分子机制,可为农作物的抗旱育种提供新型的分子标记和重要靶基因,最终服务于农业生产实践。     

植物表皮蜡质与抗旱及其分子生物学

植物表皮蜡质是覆盖在植物最外层的一类有机混合物总称,它是植物自我防护的最后一道屏障,在植物生长发育过程中起重要作用。文章就植物表皮蜡质的成分、转运、晶体的形成和发育、形态结构以及环境变化对表皮蜡质的影响,特别是在蜡质与角质蒸腾、叶片水分利用效率和产量的关系以及蜡质的生理作用和合成过程的分子生物学研究进展方面做了系统综述,并对植物蜡质研究中存在的问题做了探讨。     

植物蜡质及其与环境的关系

陆生植物的地上部分如叶、茎、花、果实等的表面覆盖着一层蜡质,它是由一系列复杂化合物组成的具有三维微结构的疏水层,在植物生长和发育过程中起着不可或缺的作用,具有很好的生物学功能。作为植物与环境的第一接触面,蜡质对外界环境因子的响应较敏感,当植物受到外界不利环境因子胁迫时,蜡质会改变自身晶体结构形态或化学组分构建防御机制以减少胁迫因子的作用,有效地协调植物与环境的关系。综述了近年来国内外关于植物蜡质的研究进展,在阐述蜡质层结构及其化学组分的基础上,着重介绍植物与环境因子的作用,包括非生物环境因子如水分、温度、光照、环境污染等以及植食性昆虫和病原菌等生物环境因子的作用。研究显示,胁迫环境下植物蜡质化学组分的变化,是由于不利环境因子的作用足以改变蜡质各产物的合成途径,从而影响蜡质产物。植物蜡质利用各种生理、化学机制对胁迫环境因子的适应以及响应,是植物适应各种生境的基础,因此通过对植物蜡质与环境关系的研究为进一步解析植物与环境关系提供证据。     

中国干旱半干旱区荒漠植物叶片(或同化枝)表皮微形态特征

为了探讨荒漠植物叶片表皮微形态结构对长期荒漠环境的适应特征及其分类学意义, 应用扫描电镜对中国干旱半干旱荒漠区28科74属117种200多个自然居群的植物叶片(或同化枝)表皮微形态进行了研究。荒漠植物叶(或同化枝)表皮的基本特征是: 表皮附属物相当丰富, 包括大量的表皮绒毛、角质膜蜡质片层或晶体颗粒、表面瘤状或疣状突起以及相对下陷且密度较低的气孔器。对表皮微形态结构及附属物组成进行对比分析, 将荒漠植物粗分为11种基本类型, 包括表皮完全被形态各异的蜡质层或表皮毛覆盖、不同形态类型的表皮毛和蜡质层结合、蜡质层与不同分布类型的气孔器或表皮毛结合, 以及各种突起的表皮细胞与蜡质层的结合等。根据抗逆所依赖的表皮及其附属物微形态结构, 将荒漠植物适应环境胁迫的叶片表皮微形态分为6种主要类型, 它们分别依赖于表皮毛、角质层蜡质、表皮凹凸结构、表面突起、混生的附属物以及上下表皮异化特征。荒漠植物叶表皮微形态结构的适应特征是通过表皮附属物(绒毛和角质膜蜡质层)与表皮结构(凹凸、乳突和气孔器)的相互协调作用, 共同抵御强光、降低叶片的蒸腾来提高植物对干旱等不利环境的抗性。该研究在一定程度上阐明了荒漠植物对逆境的适应机理及其演化趋势, 并为优良固沙植物的筛选提供了理论依据。     

小麦蜡质的逆境胁迫

蜡质是植物与外部环境的界面,对植物具有重要的生物学意义。本文综述了小麦叶表皮蜡质的物理结构、化学成分及小麦蜡质在不同逆境胁迫下的变化。     

植物表皮蜡质生物合成及调控

植物表皮蜡质,是覆盖在陆地植物地上部分表面的一层疏水性脂类物质,是植物应对外界环境变化的第一道屏障,在抑制植物水分非气孔散失及保护植物免受病虫害入侵、紫外线辐射等方面起着重要作用。综述了近年来表皮蜡质在生物合成及调控等方面的研究进展,并对研究中存在的问题及研究前景进行了展望。     

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.     

空气湿度与土壤水分胁迫对紫花苜蓿叶表皮蜡质特性的影响

选用2个抗旱性不同的紫花苜蓿品种,敖汉(强抗旱)和三得利(弱抗旱),设置空气湿度(45%-55%和75%-85%)和土壤水分胁迫(75%和35%田间持水量)处理,分析紫花苜蓿叶表皮蜡质含量、组分及晶体结构、气体交换参数、水势及脯氨酸含量的变化规律。结果表明,单独土壤水分胁迫时,紫花苜蓿叶表皮蜡质晶体结构及蜡质总量无显著变化;敖汉蜡质组分中烷类、酯类含量增加,醇类含量下降;三得利醇类含量下降,烷类、酯类含量变化不显著。低空气湿度胁迫时,两品种蜡质总量无显著变化,烷类和酯类含量显著增加,醇类含量显著下降,叶表皮片状蜡质晶体结构熔融呈弥漫性,扩大了对叶表面积的覆盖,其蒸腾速率显著低于正常湿度。复合胁迫处理时,叶表皮片状蜡质晶体结构继续呈弥漫性,烷类、酯类、未知蜡质组分含量均高于单独胁迫处理,醇类含量最低,而蜡质总量除三得利显著高于对照外,其余均无显著差异。紫花苜蓿叶表皮蜡质各组分含量(除醇类)及蜡质总量与光合速率呈显著负相关,与蒸腾速率无显著相关关系。蜡质总量与叶水势呈显著正相关。总体上,敖汉蜡质总量显著高于三得利,蜡质组分中烷类物质的增加有助于提高植株的抗旱性。在复合胁迫下,强抗旱品种主要通过气孔因素控制水分散失,而弱抗旱品种通过气孔和非气孔因素共同控制植物水分散失。     

植物角质层内外蜡质的差异及其与抗逆性的关系

植物角质层是覆盖在植物地上部分的叶、花和非木质茎等器官表面的保护层,包括角质和蜡质。其中蜡质根据分布位置不同又分为表皮蜡质和内部蜡质。大量研究表明,表皮蜡质含量和结构在植物生长发育和抗逆性申发挥着重要作用。近年来有研究发现构成蜡质的成分在内外蜡质层中的分布存在差异,角质层蜡质成分影响植物抗逆性。本文针对角质层结构和内外蜡质差异性以及角质层结构和组成与植物抗逆性之间的关系进行了综述。     

28种柳属植物的叶表皮微形态特征及其分类学意义

对28种表型相似、种间界限模糊的柳属植物在扫描电子显微镜下的叶表皮微形态特征进行观察,结果表明:柳属中有7种角质层蜡质纹饰,分别是平滑蜡质层、壳状蜡质层、痂状蜡质层、片状晶体、膜片状晶体、锥形纤维体和鳞片状纤维体,其中锥形纤维体和鳞片状纤维体为柳属所特有,而片状晶体和膜片状晶体为首次在柳属植物中发现;扫描电子显微镜下柳属植物叶表皮毛被的微观形态特征并不似其宏观形态(疏毛、绢毛、绒毛等)那样具有显著差异,微观形态主要表现为毛被密度、长短和卷曲方式(分为直,微弯曲和深度卷曲三种)的不同.研究表明叶表皮蜡质纹饰类型、气孔器的形态等微形态特征较为稳定,对柳属植物中表型相似的种类有很好的鉴定价值,但对组、亚属水平的界定作用不大;分布于寒冷地区和高海拔地区的柳属植物的叶表皮微形态特征相对多样,这可能是植物对寒冷环境的适应进化.     

Major Evolutionary Trends in Hydrogen Isotope Fractionation of Vascular Plant Leaf Waxes

Hydrogen isotopic ratios of terrestrial plant leaf waxes (δD) have been widely used for paleoclimate reconstructions. However, underlying controls for the observed large variations in leaf wax δD values in different terrestrial vascular plants are still poorly understood, hampering quantitative paleoclimate interpretation. Here we report plant leaf wax and source water δD values from 102 plant species grown in a common environment (New York Botanic Garden), chosen to represent all the major lineages of terrestrial vascular plants and multiple origins of common plant growth forms. We found that leaf wax hydrogen isotope fractionation relative to plant source water is best explained by membership in particular lineages, rather than by growth forms as previously suggested. Monocots, and in particular one clade of grasses, display consistently greater hydrogen isotopic fractionation than all other vascular plants, whereas lycopods, representing the earlier-diverging vascular plant lineage, display the smallest fractionation. Data from greenhouse experiments and field samples suggest that the changing leaf wax hydrogen isotopic fractionation in different terrestrial vascular plants may be related to different strategies in allocating photosynthetic substrates for metabolic and biosynthetic functions, and potential leaf water isotopic differences.     

Influence of atmospheric CO2 enrichment, soil N, and water stress on needle surface wax formation in Pinus palustris (Pinaceae)

Interactive effects of increasing atmospheric CO2 with resource limitations on production of surface wax in plants have not been studied. Pinus palustris seedlings were grown for 1 yr at two levels of soil N (40 or 400 kg N_ha-1_yr-1) and water stress (-0.5 or -1.5 MPa xylem pressure potential) in open-top field chambers under two levels of CO2 (365 or 720 mumol/mol). Needle surface wax content was determined at 8 mo (fall) and 12 mo (spring) and epicuticular wax morphology was examined using scanning electron microscopy (SEM) at 12 mo. Wax content expressed on both a leaf area and dry mass basis was increased due to main effects of low N and water stress. No main effects of CO2 were observed; however, a CO2 x N interaction at 12 mo indicated that under low soil N the elevated CO2 treatment had less wax (surface area or dry mass basis) compared to its ambient counterpart. Morphologically, low N needle surfaces appeared rougher compared to those of high N needles due to more extensive wax ridges. Although the main effect of water treatment on wax density was not reflected by changes in wax morphology, the CO2 x N interaction was paralleled by alterations in wax appearance. Decreases in density and less prominent epicuticular wax ridges resulting from growth under elevated CO2 and limiting N suggest that dynamics of plant/atmosphere and plant/pathogen interactions may be altered.     

The Role of Salicylic Acid Signal in Plant Growth,Development and Abiotic Stress

In nature, plants are constantly affected by adverse conditions. Unlike animals, plants can resist these adverse stresses only by insisting on their original positions. Stress can be divided into biological stress and abiotic stress, abiotic stress directly affects the growth, development and yield of plants, it spans all developmental stages from seed germination to senescence. In order to adapt to changing environment, plants have evolved well-developed mechanisms that help to perceive the stress signals and enable optimal growth response. Salicylic acid (SA) is an important endogenous signal molecule in plants, which not only regulate some plant growth and development processes, but also plays an important part in plant stress resistance. Much work about salicylic acid has been done on the immunity of plants to pathogens, and the synthesis and signal transduction of SA are clearly understood, its function in plant growth, development and abiotic stress is also well learned, we systemically summarized the multiple function of SA signal in non-pathogen-related response, such review should help us understand the common but essential function of SA signal in modulating plant growth, development and abiotic stress.     

Leaf (or assimilation branch) epidermal micromorphology of desert plant in arid and semi- arid areas of China

Aims Leaf epidermal micromorphology is an important adaptation of desert plants to arid environment. A micromorphological analysis of leaf epidermal tissue of desert plants was carried out in order to obtain qualitative and quantitative data on epidermal characteristics and to evaluate the long-term adaptive strategy of desert plants to aridity in desert conditions.
Methods The leaf (or assimilation branches) materials were sampled for more than 200 natural populations of 117 desert plant species from 74 genera and 28 families, in arid and semi-arid areas of China. The characteristics of leaf epidermal micromorphology of desert plants were then measured by scanning electron microscopy (SEM). Characteristics of epidermal cell, trichome, stomatal, cuticular wax on adaxial and abaxial surface are presented.
Important findings Leaf epidermal micromorphology of desert plants showed abundant diversity in different classification levels. The desert plants adapted to environmental stress can be divided into 11 basic morphological types according to the structure of the epidermis, and their characteristics of leaf epidermal morphology were classfied into 6 main types according to the relationships between stress resistance and structural characteristics of epidermal micromorphology and their appendages. The main epidermal appendages of desert plants (such as trichome, cuticular wax) and epidermal structures (concave-convex and papillary structure, stomata) could cooperate with each other to improve the resistance of desert plants to drought and other adverse environmental stress by resisting the strong light and reducing leaf transpiration.     

Overexpression of the ABC transporter gene <Emphasis Type="Italic">TsABCG11</Emphasis> increases cuticle lipids and abiotic stress tolerance in <Emphasis Type="Italic">Arabidopsis</Emphasis>

The cuticle, composed primarily of wax and cutin, covers most plant aerial surfaces and plays a vital role in interactions between plants and their environment. Some ATP-binding cassette G subfamily (ABCG) members are involved in cuticular lipid molecule exportation to outside in the plant surface. Thellungiella salsugineum, a relative of Arabidopsis thaliana with a heavy cuticle, has extreme stress tolerance. TsABCG11, an ABCG transporter was cloned (GenBank accession number JQ389853), and its structure was studied. qRT-PCR showed that TsABCG11 expression varied in different organs of T. salsugineum and was upregulated under ABA, NaCl, drought and cold conditions. The rosette leaves from 4-week-old TsABCG11 overexpressed (OE) Arabidopsis plants displayed lower rates of water loss and decreased chlorophyll-extracted rates compared to wild-type plants. TsABCG11-OE plants also exhibited significantly increased total cuticular wax and cutin monomer amounts, mainly due to prominent changes in the C₂₉, C₃₁, and C₃₃ alkanes in the wax and C18:2 dioic in cutin monomers, respectively. TsABCG11-OE seedlings exhibit lower root growth inhibition under 100 mM of NaCl or 1 µM of ABA than the wild type. Four-week-old TsABCG11-OE plants exhibited higher photosynthetic rates and water-use efficiency under cold stress (4 °C) than control plants. These results indicate that TsABCG11 plays an important role in cuticle lipid exportation and is involved in abiotic stresses, probably having a close relationship with extreme stress tolerance in T. salsugineum.     

Plant response to drought stress simulated by ABA application: Changes in chemical composition of cuticular waxes

Plant cuticles form the interface between epidermal plant cells and the atmosphere. The cuticle creates an effective barrier against water loss, bacterial and fungal infection and also protects plant tissue from UV radiation. It is composed of the cutin matrix and embedded soluble lipids also called waxes. Chemical composition of cuticular waxes and physiological properties of cuticles are affected by internal regulatory mechanisms and environmental conditions (e.g. drought, light, and humidity). Here, we tested the effect of drought stress simulation by the exogenous application of abscisic acid (ABA) on cuticular wax amount and composition. ABA-treated plants and control plants differed in total aboveground biomass, leaf area, stomatal density and aperture, and carbon isotope composition. They did not differ in total wax amount per area but there were peculiar differences in the abundance of particular components. ABA-treated plants contained significantly higher proportions of aliphatic components characterized by chain length larger than C₂₆, compared to control plants. This trend was consistent both between and within different functional groups of wax components. This can lead to a higher hydrophobicity of the cuticular transpiration barrier and thus decrease cuticular water loss in ABA-treated plants. At both ABA-treated and control plants alcohols with chain length C₂₄ and C₂₆ were predominant. Such a shift towards wax compounds having a higher average chain length under drought conditions can be interpreted as an adaptive response of plants towards drought stress.