甘蓝型油菜叶表皮蜡质组分及结构与菌核病抗性关系

本试验选用6个抗病性不同的甘蓝型油莱品种,研究其叶表皮蜡质组成及结构与菌核病抗性的关系。结果表明,抗病品种在去除叶表皮蜡质后病情指数显著增加;感病品种无显著变化。不同抗性品种（系）间除酯类组分含量无显著差异外,其余蜡质组分含量差异显著。相关分析表明,蜡质组分中酯类含量与病情指数呈显著负相关关系,醇类、酮类含量与病情指数呈显著正相关,其余组分和蜡质总量与病情指数无显著相关关系。抗性品种叶表皮蜡质中烷类及酯类所占比重较高,而易感品种酮类比重较高。扫描电镜结果显示,抗病品种（系）的蜡质晶体主要为颗粒状、杆状、丝状;而感病品种（系）的蜡质晶体中不规则片状晶体所占比例较大。这些结果说明油菜叶表皮蜡质的组分及结构可能是抗病品种抵抗和延迟病原菌侵入的机制之一。     

核盘菌侵染对拟南芥表皮蜡质结构及化学组成的影响

以野生型拟南芥与蜡质突变体cer1、cer4为试验材料,通过研究核盘菌胁迫对拟南芥茎表皮蜡质结构及组分含量的影响,揭示核盘菌侵染与表皮蜡质的关系。扫描电镜结果显示,野生型拟南芥蜡质晶体以垂直于表面的杆状、块状结构为主;突变体cer1晶体类型以水平的松针状、块状结构为主;突变体cer4蜡质晶体以垂直片层结构为主。核盘菌胁迫下,拟南芥蜡质晶体结构及分布形态发生变化。蜡质层结构在核盘菌胁迫下表现为:杆状、松针状蜡质晶体减少—蜡质晶体熔融—表皮"囊状凸起"—表皮膜层破裂。这些结构变化有利于病菌突破角质层屏障而侵入到植株体内。色质谱分析结果显示:与野生型相比,cer1突变体烷、次级醇、酮类显著减少;cer4突变体表现为一级醇含量减少。接种核盘菌后,野生型拟南芥与蜡质突变体一级醇类显著增加(cer1增加不显著);烷类、次级醇类、酮类含量与蜡质总量均显著减少,表明蜡质前体物质在受到核盘菌胁迫后更多地通过酰基还原途径生成一级醇,从而减少了由脱羰基途径所生成的蜡质组分。核盘菌通过改变表皮蜡质晶体结构与化学组分分泌量来促进侵染。     

高羊茅叶片表皮蜡质含量与其抗旱性的关系

以14个高羊茅品种为试验材料,在田间试验中对干旱高温胁迫下的叶片表皮蜡质含量、净光合速率、蒸腾速率、气孔导度、胞间CO2浓度等生理指标测定分析。结果表明,干热胁迫下高羊茅品种间的叶片表皮蜡质含量和水分利用效率均存在极显著差异(P<0.01);叶片蜡质含量与综合抗旱性和水分利用效率的等级相关系数分别为0.78(P<0.01)和0.68(P<0.01);蜡质含量越高的品种,其叶片气孔导度和胞间CO2浓度越低,水分利用效率越高,但所有品种的水分利用效率绝对值都较低。研究发现,在干热胁迫时,高羊茅叶片表皮蜡质可通过对气孔导度的调节来减少气孔蒸腾,提高水分利用效率,最终提高其抗旱性;表皮蜡质含量可以作为高羊茅品种抗旱性鉴定的一个新指标。     

宁夏枸杞果皮蜡质微形态及蜡质组分研究北大核心CSCD

果实制干是宁夏枸杞炮制的主要过程,宁夏枸杞不同品种在制干特性方面存在一定的差异。该研究以宁夏枸杞栽培中制干差异较大的品种‘宁杞1号’(易制干)和‘宁杞5号’(不易制干)果实为材料,采用扫描电镜技术和GC-MS技术对2个宁夏枸杞品种不同发育时期(青果期、色变期、成熟期)的果实果皮结构以及果皮蜡质微形态、含量和组分进行了观测,从果皮蜡质微形态及组分的积累变化初步揭示枸杞果皮蜡质的积累规律,以明确不同枸杞品种果皮蜡质组分差异,为不同品种适宜促干剂的筛选以及促干剂的合理使用提供理论依据。结果表明:(1)‘宁杞1号’和‘宁杞5号’枸杞表皮细胞外侧细胞壁均呈现脊状突起的结构,在果实青果期脊状突起不连续,脊和脊之间排列紧密;随着发育时期的延后,脊状突起的连续性逐渐增强,且脊与脊之间的间距逐渐变宽,蜡质呈膜状覆盖于凸起的脊和两脊之间的沟内。(2)‘宁杞1号’和‘宁杞5号’两个枸杞品种在果实发育过程中,果皮单位面积蜡质含量均呈先下降后上升的趋势,在成熟期单位面积蜡质含量最高,且‘宁杞1号’3个发育时期果皮单位面积蜡质含量均高于同期‘宁杞5号’,呈现出与扫描电镜观察到的蜡质分布基本一致的变化趋势。(3)2个品种3个时期的果实表皮蜡质组分均由烷烃类、酮类、醇类、酸类、醛类、酯类和碘代烷烃类组成,两品种青果期和色变期果皮蜡质组分相同,主要由烷烃类、醇类和碘代烷烃类组成,它们成熟期的果皮蜡质组分主要由烷烃类、醇类和酯类组成。(4)主成分分析结果显示,3个生育时期‘宁杞5号’果皮蜡质组分多为烷烃类物质,较‘宁杞1号’更利于阻挡果实水分的散失。研究发现,影响枸杞果实制干的原因在于枸杞果皮蜡质含量、结构和蜡质组分,烷烃类组分能够有效阻止果实体内水分的散失,‘宁杞5号’果皮蜡质中烷烃组分含量更高,果皮保水性更强,致果实不易制干。     

UV-B辐射增强对拟南芥表皮蜡质的影响

以野生型拟南芥、蜡质不同程度缺失突变体CER1、CER3、CER4、CER6、CER10、CER20及KCS1为试验材料,通过施加50μW/cm2、长达10 d的UV-B辐射,研究了拟南芥表皮蜡质晶体结构、组分及蜡质基因对UV-B辐射的响应机制。结果表明:UVB辐射增强改变了拟南芥表皮蜡质晶体结构,表皮蜡质松针状(CER1)、柱状、杆状(CER3、CER10与KCS1)晶体结构显著减少,球状蜡质晶体类型出现在CER6表面,无规则片状、膜状结构覆盖在KCS1与CER10茎表面。野生型拟南芥蜡质晶体结构类型无明显变化,但在部分区域积累了大量水平杆状、管状结构,增加了蜡质层厚度。UV-B辐射增强也改变了拟南芥表皮蜡质组分的分泌量。野生型在UV-B处理后一级醇、酸、醛含量显著上升,烷、次级醇及酮含量显著下降,蜡质总量增加不显著。一级醇含量的增加及酮和次级醇含量的减少在拟南芥各材料响应UV-B辐射中具有普遍性。UV-B辐射增强诱导了野生型CER3、CER4、KCS1基因表达的上调,其中CER4大量表达,促进了蜡质组分中一级醇、酸和醛含量的积累;CER1在UV-B处理后表达量下调,可能导致烷合成下游分支途径相关产物(烷类、次级醇及酮类)的减少。WIN1表达量的下调对蜡质总量没有显著影响。UV-B辐射增强使蜡质前体从烷合成分支途径更多地转向一级醇分支途径。     

小麦品系CP98(11)不同器官表皮蜡质组分和蜡质晶体结构的差异分析

为明确小麦不同器官表皮蜡质晶体结构和蜡质组分的差异,该研究以小麦品系CP98(11)为材料,在小麦扬花期分别取小麦的旗叶、叶鞘、穗下茎、花药和颖壳,利用气相色谱技术对各器官表皮蜡质组分进行鉴定,并通过扫描电镜观察其蜡质晶体结构。结果表明:(1)小麦不同器官的蜡质成分共鉴定出30种,主要为初级醇、二酮、烷烃、脂肪醛、脂肪酸、酯。(2)叶鞘、穗下茎、颖壳的蜡质中二酮含量最高,分别占蜡质总量的78.96%、67.03%和68.6%;花药的蜡质中烷烃含量最高(75.82%);旗叶的蜡质中初级醇含量最高(45.91%),其次为烷烃33.19%。(3)扫描电镜观察显示,旗叶正面的蜡质晶体呈片状结构,旗叶反面和颖壳的蜡质晶体结构呈片状与柱状混合的结构,花药的蜡质呈明显的波浪状结构,穗下茎和叶鞘的蜡质晶体呈柱状结构。     

高粱表皮蜡质缺失突变体sb1抗旱生理生化分析

植物抗旱性与体表蜡质积累有关，高粱是抗旱性极强的作物，茎秆和叶片表面覆盖一层厚厚的蜡质，开展高粱体表蜡质层对高粱抗旱能力的研究，为高粱抗旱品种选育和抗旱分子机制提供理论依据。以高粱品种BTx623和表皮蜡质缺失突变体sb1为试验材料，统计农艺性状以及扫描电镜观察叶片表面蜡质形态结构；分析表皮蜡质的有无对离体叶片失水速率和叶绿素浸提率的影响；检测在干旱胁迫处理下植株的表型变化以及超氧化物歧化酶（SOD）、过氧化物酶（POD）的活性变化。结果表明，与对照BTx623相比，表皮蜡质缺失突变体sb1株高明显降低、抽穗开花期延后1周、叶片远轴面片状蜡质缺失，其他农艺性状变化不明显。突变体sb1离体叶片失水速率和叶绿素浸提率显著高于对照。在干旱胁迫处理条件下，随着干旱胁迫时间的增长，对照BTx623和突变体sb1的SOD和POD活性均增强，但对照SOD和POD活性均显著高于突变体sb1，且干旱处理96 h和复水后突变体sb1都造成叶片大面积干枯。突变体sb1表皮蜡质缺失后叶片渗透性增强，清除活性氧的能力减弱，抗旱性降低。     

不同抗旱性冬小麦幼苗根系对水分胁迫的反应

抗旱性不同的小麦根系含水量、水势、渗透势均随水分胁迫强度增加而逐渐下降。其中以水势变化最为灵敏。恢复正常供水72h后,三项指标均有不同程度的回升,抗旱品种恢复能力强。根系渗透调节能力随胁迫强度的加剧而提高,抗旱品种渗透调节的效果好于敏感品种。随着胁迫强度的增加,根中ATP相对含量减少,恢复正常供水72h后,含量可部分恢复,恢复能力与品种的抗旱性一致。     

不同抗旱性冬小冬幼苗根系对小分胁迫的反应

抗旱性不同的小麦根系含水量、水势、渗透势均随水分胁迫强度增加而逐渐下降。其中以水势变化最为灵敏。恢复正常供水７２ｈ后,三项指标均有不同程度的回升,抗旱品种恢复能力强。根系渗透调节能力随胁迫强度的加剧而提高,抗旱品种渗透调节的效果好于敏感品种。随着胁迫强度的增加,根中ＡＴＰ相对含量减少,恢复正常供水７２ｈ后,含量可部分恢复,恢复能力与品种的抗旱性一致。     

干旱胁迫下不同抗旱性小麦叶片内源激素含量的变化与抗旱力强弱的关系

以抗旱性强的小麦品种昌乐5号和抗旱性弱的鲁麦5号的幼苗为材料,研究了随着干旱胁迫的加剧,小麦叶片相对含水量、气孔导度和内源激素水平的变化．结果表明,叶片的气孔导度和细胞激动素与脱落酸含量的比值（CTKs/ABA）呈较强的线性正相关关系而与叶片RWC的相关关系较弱,说明气孔导度受CTKs/ABA调控,而不是受叶片RWC的调控．在不同强度的干旱胁迫下我们可以利用CTKs/ABA的变化判断品种抗旱性大小：①抗旱力强的小麦品种叶片的平均CTKs/ABA值较高,而弱者较低,说明强者内源促进型激素含量相对较高,从而在干旱胁迫下保持较高的生活力．②抗旱力强的小麦品种叶片CTKs/ABA与干旱处理天数呈二次负相关关系,而弱者是线性负相关关系,说明强者具有较强的抗逆缓冲能力．③抗旱力强的小麦品种成熟叶（第二叶）CTKs／ABA下降快,新叶（第三叶）下降慢,而弱者反之,说明抗旱性强者可能存在着较强的从成熟叶向新叶的物质运输从而具有自我保护性调节机制．     

Leaf cuticular waxes and physiological parameters in alfalfa leaves as influenced by drought

Drought significantly constrains higher yield of alfalfa (Medicago sativa L.) in arid and semiarid areas all over the world. This study evaluated the responses of leaf cuticular wax constituents to drought treatment and their relations to gas-exchange indexes across six alfalfa cultivars widely grown in China. Water deficit was imposed by withholding water for 12 d during branching stage. Cuticular waxes on alfalfa leaves were dominated by primary alcohols (41.7?C54.2%), alkanes (13.2?C26.9%) and terpenes (17.5?C28.9%), with small amount of aldehydes (1.4?C3.4%) and unknown constituents (4.5?C18.4%). Compared to total wax contents, the wax constituents were more sensitive to drought treatment. Drought decreased the contents of primary alcohol and increased alkanes in all cultivars. Alkane homologs, C25, C27, and C29, were all negatively correlated with photosynthetic rate, transpiration rate, stomatal conductance, and leaf water potential. Under drought conditions, both stomatal and nonstomatal factors were involved in controlling water loss from alfalfa leaves. No direct relationship was observed between wax contents and drought resistance among alfalfa cultivars. An increase in alkane content might be more important in improving drought tolerance of alfalfa under water deficit, which might be used as an index for selecting and breeding drought resistant cultivars of alfalfa.     

Leaf cuticular waxes of potted rose cultivars as affected by plant development, drought and paclobutrazol treatments

The present work was carried out to evaluate how plant growth and cultural practices influence the amount and composition of cuticular waxes on leaves of rose cultivars. The total amount of cuticular wax per leaf area was higher for rose cultivar Apollo Parade than for Charming Parade. Both cultivars had waxes dominated by alkanes, with the major alkanes being the C₃₁ and C₃₃ homologues. Primary alcohols were the next most abundant constituent class, with C₂₆ as the dominant homologue. Compared with Charming Parade, Apollo Parade had higher proportions of its total wax load as primary alcohols but lower acids and aldehydes. The proportion of alkanes in the total load on these cultivars was similar. Commercially produced roses are routinely treated with paclobutrazol (PBZ) to retard growth. PBZ treatments caused a 10% increase in total wax load and changes in the proportions of certain wax constituents within 11 days of application. Notable was an increase in the total proportion of acids in the total load 25 days after PBZ application, primarily because of increased C₂₈ acids. An alternative method of retarding plant growth is production of roses under limited water availability. When Apollo Parade roses experienced periods of moderate drought stress during production, the wax load per leaf area increased 14 and 8% above control levels at 24 and 38 days after imposition of drought, respectively. Drought caused similar changes in the proportions of individual wax constituents as did PBZ application.     

Diversity of cuticular wax among Salix species and Populus species hybrids

The leaf cuticular waxes of three Salix species and two Populus species hybrids, selected for their ability to produce high amounts of biomass, were characterized. Samples were extracted in CH(2)Cl(2) three times over the growing season. Low kV SEM was utilized to observe differences in the ultrastructure of leaf surfaces from each clone. Homologous series of wax components were classified into organic groups, and the variation in wax components due to clone, sample time, and their interaction was identified. All Salix species and Populus species hybrids showed differences in total wax load at each sampling period, whereas the pattern of wax deposition over time differed only between the Salix species. A strong positive relationship was identified between the entire homologous series of alcohols and total wax load in all clones. Similarly strong relationships were observed between fatty acids and total wax load as well as fatty acids and alcohols in two Salix species and one Populus species hybrid. One Salix species, S. dasyclados, also displayed a strong positive relationship between alcohols and alkanes. These data indicate that species grown under the same environmental conditions produce measurably different cuticular waxes and that regulation of wax production appears to be different in each species. The important roles cuticular waxes play in drought tolerance, pest, and pathogen resistance, as well as the ease of wax extraction and analysis, strongly suggest that the characteristics of the cuticular wax may prove to be useful selectable traits in a breeding program.     

Changes in leaf cuticular waxes of sesame (Sesamum indicum L.) plants exposed to water deficit

Sesame (Sesamum indicum L.) is one of the most important oilseed crops, having seeds and oil that are highly valued as a traditional health food. The objective of this study was to evaluate leaf cuticular wax constituents across a diverse selection of sesame cultivars, and the responses of these waxes to drought-induced wilting. Water-deficit was imposed on 18 sesame cultivars by withholding irrigation for 15d during the post-flowering stage, and the effect on seed yield and leaf waxes compared with a well-watered control. Leaf cuticular waxes were dominated by alkanes (59% of total wax), with aldehydes being the next-most abundant class. Compared to well-irrigated plants, drought treatment caused an increase in wax amount on most cultivars, with only three cultivars having a notable reduction. When expressed as an average across all cultivars, drought treatment caused a 30% increase in total wax amount, with a 34% increase in total alkanes, a 13% increase in aldehydes, and a 28% increase in the total of unknowns. In all cultivars, the major alkane constituents were the C27, C29, C31, C33, and C35 homologs, whereas the major aldehydes were the C30, C32, and C34 homologs, and drought exposure had only minor effects on the chain length distribution within these and other wax classes. Drought treatments caused a large decrease in seed yield per plant, but did not affect the mean weight of individual seeds, showing that sesame responds to post-flowering drought by reducing seed numbers, but not seed size. Seed yield was inversely correlated with the total wax amount (-0.466*), indicating that drought induction of leaf wax deposition does not contribute directly to seed set. Further studies are needed to elucidate the ecological role for induction of the alkane metabolic pathway by drought in regulating sesame plant survival and seed development in water-limiting environments.     

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.     

Ontogenetic variation in chemical and physical characteristics of adaxial apple leaf surfaces

The reaction of plants to environmental factors often varies with developmental stage. It was hypothesized, that also the cuticle, the outer surface layer of plants is modified during ontogenesis. Apple plantlets, cv. Golden Delicious, were grown under controlled conditions avoiding biotic and abiotic stress factors. The cuticular wax surface of adaxial apple leaves was analyzed for its chemical composition as well as for its micromorphology and hydrophobicity just after unfolding of leaves ending in the seventh leaf insertion. The outer surface of apple leaves was formed by a thin amorphous layer of epicuticular waxes. Epidermal cells of young leaves exhibited a distinctive curvature of the periclinal cell walls resulting in an undulated surface of the cuticle including pronounced lamellae, with the highest density at the centre of cells. As epidermal cells expanded during ontogenesis, the upper surface showed only minor surface sculpturing and a decrease in lamellae. With increasing leaf age the hydrophobicity of adaxial leaf side decreased significantly indicated by a decrease in contact angle. Extracted from plants, the amount of apolar cuticular wax per area unit ranged from only 0.9 microgcm(-2) for the oldest studied leaf to 1.5 microgcm(-2) for the youngest studied leaf. Differences in the total amount of cuticular waxes per leaf were not significant for older leaves. For young leaves, triterpenes (ursolic acid and oleanolic acid), esters and alcohols were the main wax components. During ontogenesis, the proportion of triterpenes in total mass of apolar waxes decreased from 32% (leaf 1) to 13% (leaf 7); absolute amounts decreased by more than 50%. The proportion of wax alcohols and esters, and alkanes to a lesser degree, increased with leaf age, whereas the proportion of acids decreased. The epicuticular wax layer also contained alpha-tocopherol described for the first time to be present also in the epicuticular wax. The modifications in the chemical composition of cuticular waxes are discussed in relation to the varying physical characteristics of the cuticle during ontogenesis of apple leaves.     

Crystal structures and chemical composition of leaf surface wax depositions on the desert moss Syntrichia caninervis

The structures and composition of cuticular waxes deposited on the leaves of a typical desert moss, Syntrichia caninervis, were investigated. The wax crystals deposited on leaves shifted with leaf aging. The results of chemical analysis showed the main chemical components of the moss wax were fatty acids, alcohols and alkanes. Leaf aging increased the content of cuticular wax and the percentage of very long chain components, from 1150 μg g⁻¹ DW and 13.6% in younger leaves to 2640 μg g⁻¹ DW and 37.2% in aged leaves, respectively. Dehydration/hydration also augmented the wax content by 35.17% in juvenile leaves and by 1900% in lab-cultivated leaves after three-cycle treatments. Synthesis of hexadecanoic acid and tetracosane were predicted to be the first step of wax accumulation. The responses of cuticular waxes in crystal structure and chemical composition were recommended as a biomonitor for assessing the shift of ecological and environmental quality.     

A morpho-physiological approach differentiates bread wheat cultivars of contrasting tolerance under cyclic water stress

Leaf micromorphological traits and some physiological parameters with potential relevance to drought tolerance mechanisms were investigated in four selected winter wheat varieties. Plants were subjected to two cycles of drought treatment at anthesis. Yield components confirmed contrasting drought-sensitive and -tolerant behavior of the genotypes. Drought tolerance was associated with small flag leaf surfaces and less frequent occurrence of stomata. Substantial variation of leaf cuticular thickness was found among the cultivars. Thin cuticle coincided with drought sensitivity and correlated with a high rate of dark-adapted water loss from leaves. Unlike in Arabidopsis, thickening of the cuticular matrix in response to water deprivation did not occur. Water stress induced epicuticular wax crystal depositions preferentially on the abaxial leaf surfaces. According to microscopy and electrolyte leakage measurements from leaf tissues, membrane integrity was lost earlier or to a higher extent in sensitive than in tolerant genotypes. Cellular damage and a decline of relative water content of leaves in sensitive cultivars became distinctive during the second cycle of water deprivation. Our results indicate strong variation of traits with potential contribution to the complex phenotype of drought tolerance in wheat genotypes. The maintained membrane integrity and relative water content values during repeated water limited periods were found to correlate with drought tolerance in the selection of cultivars investigated.