借助细胞晶体鉴别药用植物

在植物的细胞中,最常见是草酸钙结晶,它们是植物的一种自我解毒的过程,即对植物有毒害作用的多量草酸钙离子中和,它对植物的生长发育无明显的作用,通常认为这是植物新陈代谢产生的一种废弃物。草酸钙为无色透明的结晶,并形成不同的形态,其中有簇晶（呈霰星状）、针晶（针状）、方晶（呈斜方形、菱形、长方形）、砂晶（细小的三角形或箭头状）,广泛分布于植物各器官中。一般在一种植物中只能见到一种形态,极少数情况下也有二种或三种形态。在不同的植物中,即使晶体的形态特征相同,而其大小和数量等也不完全相同。另外,即使亲缘关…     

江西产酸模属植物叶片比较解剖学研究

对江西产酸模属(Rumex L.)7种植物进行了叶形态结构的显微观察.结果表明:(1)叶片上表皮均有气孔器分布,表皮上普遍具有腺体和异细胞,叶肉中均含有晶体细胞,晶体类型为簇晶;(2)下表皮细胞特征及气孔类型、叶肉细胞的分化及排列方式、中脉维管束数目等特征具有明显的种间差异,可以作为属下种级鉴定指标;(3)小酸模在气孔类型、栅栏组织和海绵组织排列紧密程度、中脉形状及维管束数量等特征上与同属其它种类具有显著的差异,形态解剖学证据支持小酸模亚属(Subgen.Acetosella(Meisn.)Rech.)的成立;(4)根据酸模属植物气孔器类型的演化阶段,并结合孢粉学、形态学等证据,认为酸模属处于蓼科植物系统演化的较低或中等地位.通过对酸模属植物叶形态结构的比较观察,为探讨该属的系统演化关系及属下分类提供叶解剖学证据.     

五种C4荒漠植物光合器官中含晶细胞的比较分析

为了探讨荒漠植物适应干旱环境的机理,选择光合器官发生很大变化的5种C4荒漠植物进行了解剖结构的对比研究。结果表明,这5种植物中含晶细胞的数量、大小、形态和分布位置等存在差异。白梭梭（Haloxylon persicum）和梭梭（H.ammodendron）的同化枝普遍具有含晶细胞;沙拐枣（Calligonum mongolicum）的含晶细胞很少,一般只分布在贮水组织或靠近栅栏组织处;木本猪毛菜（Salsola arbuscula）的含晶细胞也不多,主要分布在栅栏组织和表皮细胞之间;猪毛菜（S.collina）的含晶细胞更少,仅在贮水组织中偶尔可见晶簇。比较梭梭、白梭梭和沙拐枣同化枝不同部位的解剖结构发现,梭梭同化枝基部含晶细胞最多,中部次之,顶部最少;白梭梭同化枝项部的含晶细胞数量较多,中部及基部较少;沙拐枣同化枝顶部与基部的粘液细胞较多,中部较少,基部几乎没有栅栏组织,而其维管组织较为发达。综合晶体的酸碱溶解性及硝酸银组化分析结果,并参照能谱仪的分析结果得知,梭梭、白梭梭、沙拐枣和木本猪毛菜的叶片或同化枝中所含晶体的主要成分为草酸钙。通过比较解剖结构发现,梭梭和白梭梭的同化枝中含晶细胞最多,其它3种植物的同化器官中含晶细胞较少,而沙拐枣同化枝中有粘液细胞存在。     

五种C4荒漠植物光合器官中含晶细胞的比较分析

 为了探讨荒漠植物适应干旱环境的机理, 选择光合器官发生很大变化的5种C₄荒漠植物进行了解剖结构的对比研究。结果表明, 这5种 植物中含晶细胞的数量、大小、形态和分布位置等存在差异。白梭梭(Haloxylon persicum)和梭梭(H. ammodendron)的同化枝普遍具有含晶细 胞; 沙拐枣(Calligonum mongolicum)的含晶细胞很少, 一般只分布在贮水组织或靠近栅栏组织处; 木本猪毛菜(Salsola arbuscula)的含晶细 胞也不多, 主要分布在栅栏组织和表皮细胞之间; 猪毛菜(S. collina)的含晶细胞更少, 仅在贮水组织中偶尔可见晶簇。比较梭梭、白梭梭和 沙拐枣同化枝不同部位的解剖结构发现, 梭梭同化枝基部含晶细胞最多, 中部次之, 顶部最少; 白梭梭同化枝顶部的含晶细胞数量较多, 中部 及基部较少; 沙拐枣同化枝顶部与基部的粘液细胞较多, 中部较少, 基部几乎没有栅栏组织, 而其维管组织较为发达。综合晶体的酸碱溶解性 及硝酸银组化分析结果, 并参照能谱仪的分析结果得知, 梭梭、白梭梭、沙拐枣和木本猪毛菜的叶片或同化枝中所含晶体的主要成分为草酸钙 。通过比较解剖结构发现, 梭梭和白梭梭的同化枝中含晶细胞最多, 其它3种植物的同化器官中含晶细胞较少, 而沙拐枣同化枝中有粘液细胞存 在。     

番荔枝科植物叶的比较解剖学

利用扫描电镜、叶表皮离析法和石蜡切片法研究了番荔枝科93种2变种植物叶片的形态结构.结果表明番荔枝科植物叶片形态结构具有较大相似性,如叶表面均具有表皮毛,表皮细胞具有晶体,气孔器为平列型,具2～6个副卫细胞,仅分布在远轴面,普遍具有败育气孔器,叶肉组织中普遍含有油细胞等,但表皮毛的类型,表皮细胞的形状,表皮细胞内晶体的类型和形态,叶肉组织的结构具有明显的属间和种间差异.     

杨柳科7种植物次生韧皮部的比较研究

本文研究和比较了杨柳科２属７种植物次生韧皮部解剖结构。结果表明：１杨属和柳属植物在次生韧皮部解剖上有某些共同特征：次生韧皮部具有明显分层现象;韧皮纤维和含晶细胞与筛管分子,伴胞和韧皮薄壁组织细胞呈切向带相间排列;筛管分子均为复筛板,端壁倾斜平均含量有７－８个筛域。２两属植物在射线和晶体类型上有明显区别：柳属植物次生韧皮部无石细胞;杨属植物不具功能韧皮中含有石细胞。３两属植物均有一些较为原始的韧皮剖     

杨柳科(Salicaceae)7种植物次生韧皮部的比较研究

本文研究和比较了杨柳科2属7种植物次生韧皮部解剖结构。结果表明:(1)杨属和柳属植物在次生初皮部解剖上有某些共同特征:次生韧皮部具有明显分层现象;韧皮纤维和含晶细胞与筛管分子、伴胞和韧皮薄壁组织细胞是切向带相间排列;筛管分子均为复筛板,端壁倾斜平均含有7-8个筛域。(2)两属植物在射线和晶体类型上有明显区别:柳属植物次生韧皮部无石细胞;杨属植物不具功能韧皮部中含有石细胞。(3)两属植物均有一些较为原始的韧皮部解剖特征。     

暗罗属植物叶的比较解剖学研究

利用扫描电镜技术、叶片叶片离析方法和石蜡切片法对暗罗属12种植物叶和形态结构进行了比较研究。结果表明,叶表皮细胞形状、气孔器形态、表皮毛类型、表皮细胞中晶体类型、叶肉中油细胞分布位置、栅栏组织和海绵组织厚度的比值,以及主脉维管组织的结构特征等具有明显的种间差异。可以利用这些叶的解剖特征将暗罗属植物相互区别开来。     

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.     

草酸钙结晶在几种凤仙花属植物中的特征及其分类学意义

首次研究了7种凤仙花属(ImpatiensL.)植物茎的解剖学及细胞组织中草酸钙结晶的特征.结果表明,7种凤仙花属植物茎的解剖学结构非常近似,而茎中草酸钙结晶特征则差异显著,7种凤仙花属植物茎中均有草酸钙针晶,根据草酸钙结晶形态特征的不同,将针晶分为3种类型,即针晶束、针晶簇和散针晶.其中,黄金凤、长角凤仙花、锐齿凤仙花和红纹凤仙花有针晶束分布,而湖北凤仙花、紫花黄金凤和窄萼凤仙花则无针晶束分布,只有针晶簇或散针晶分布;此外,针晶的形态、长度、排列方式及丰富程度等在不同的物种中亦有差异.草酸钙结晶特征对凤仙花属植物的分类具有一定的科学意义.     

Calcium oxalate crystals in plants

Calcium (Ca) oxalate crystals occur in many plant species and in most organs and tissues. They generally form within cells although extracellular crystals have been reported. The crystal cells or idioblasts display ultrastructural modifications which are related to crystal precipitation. Crystal formation is usually associated with membranes, chambers, or inclusions found within the cell vacuole(s). Tubules, modified plastids and enlarged nuclei also have been reported in crystal idioblasts. The Ca oxalate crystals consist of either the monohydrate whewellite form, or the dihydrate weddellite form. A number of techniques exist for the identification of calcium oxalate. X-ray diffraction, Raman microprobe analysis and infrared spectroscopy are the most accurate. Many plant crystals assumed to be Ca oxalate have never been positively identified as such. In some instances, crystals have been classified as whewellite or weddellite solely on the basis of their shape. Certain evidence indicates that crystal shape may be independent of hydration form of Ca oxalate and that the vacuole crystal chamber membranes may act to mold crystal shape; however, the actual mechanism controlling shape is unknown. Oxalic acid is formed via several major pathways. In plants, glycolate can be converted to oxalic acid. The oxidation occurs in two steps with glyoxylic acid as an intermediate and glycolic acid oxidase as the enzyme. Glyoxylic acid may be derived from enzymatic cleavage of isocitric acid. Oxaloacetate also can be split to form oxalate and acetate. Another significant precursor of oxalate in plants is L-ascorbic acid. The intermediate steps in the conversion of L-ascorbic acid to oxalate are not well defined. Oxalic acid formation in animals occurs by similar pathways and Ca oxalate crystals may be produced under certain conditions. Various functions have been attributed to plant crystal idioblasts and crystals. There is evidence that oxalate synthesis is related to ionic balance. Plant crystals thus may be a manifestation of an effort to maintain an ionic equilibrium. In many plants oxalate is metabolized very slowly or not at all and is considered to be an end product of metabolism. Plant crystal idioblasts may function as a means of removing the oxalate which may otherwise accumulate in toxic quantities. Idioblast formation is dependent on the availability of both Ca and oxalate. Under Ca stress conditions, however, crystals may be reabsorbed indicating a storage function for the idioblasts for Ca. In addition, it has been suggested that the crystals serve purely as structural supports or as a protective device against foraging animals. The purpose of this review is to present an overview of plant crystal idioblasts and Ca oxalate crystals and to include the most recent literature.     

Isolation of Medicago truncatula mutants defective in calcium oxalate crystal formation

Plants accumulate crystals of calcium oxalate in a variety of shapes, sizes, amounts, and spatial locations. How and why many plants form crystals of calcium oxalate remain largely unknown. To gain insight into the regulatory mechanisms of crystal formation and function, we have initiated a mutant screen to identify the genetic determinants. Leaves from a chemically mutagenized Medicago truncatula population were visually screened for alterations in calcium oxalate crystal formation. Seven different classes of calcium oxalate defective mutants were identified that exhibited alterations in crystal nucleation, morphology, distribution and/or amount. Genetic analysis suggested that crystal formation is a complex process involving more than seven loci. Phenotypic analysis of a mutant that lacks crystals, cod 5, did not reveal any difference in plant growth and development compared with controls. This finding brings into question the hypothesized roles of calcium oxalate formation in supporting tissue structure and in regulating excess tissue calcium.     

香樟叶肉含晶细胞季节变化研究

为探讨香樟(Cinnamomum camphora)叶肉含晶细胞超微结构的季节变化,阐明香樟叶肉中草酸钙晶体在春夏秋冬的变化规律。该研究以多年生香樟(C. camphora)叶片为材料,分别于春夏秋冬四个季节露地取样,制作超薄切片,用透射电子显微镜(TEM)观察叶肉含晶细胞超微结构的变化。结果表明:春季时香樟叶肉中只有少数细胞有草酸钙晶体,数量较少,晶体结构多为柱状晶、方晶; 夏季时香樟叶肉细胞中随机分布于液泡的草酸钙晶体明显比春季的数量多、体积大、形态丰富,晶体多为柱状晶、方晶、针晶、簇晶; 秋季时香樟叶肉细胞草酸钙晶体和夏季的类似,数量较多,形态多样,以方晶和柱状晶针晶为主,伴有晶簇; 冬季时香樟叶肉含晶细胞晶体形态为柱状晶、方晶、针晶,数量比夏季和秋季的数量略有减少。该研究结果表明在一年四季中香樟叶肉细胞液泡中均有草酸钙晶体结构存在。     

Morphologies and elemental compositions of calcium crystals in phyllodes and branchlets of Acacia robeorum (Leguminosae: Mimosoideae)

Background and Aims

Formation of calcium oxalate crystals is common in the plant kingdom, but biogenic formation of calcium sulfate crystals in plants is rare. We investigated the morphologies and elemental compositions of crystals found in phyllodes and branchlets of Acacia robeorum, a desert shrub of north-western Australia.

Methods

Morphologies of crystals in phyllodes and branchlets of A. robeorum were studied using scanning electron microscopy (SEM), and elemental compositions of the crystals were identified by energy-dispersive X-ray spectroscopy. Distributional patterns of the crystals were studied using optical microscopy together with SEM.

Key Results

According to the elemental compositions, the crystals were classified into three groups: (1) calcium oxalate; (2) calcium sulfate, which is a possible mixture of calcium sulfate and calcium oxalate with calcium sulfate being the major component; and (3) calcium sulfate · magnesium oxalate, presumably mixtures of calcium sulfate, calcium oxalate, magnesium oxalate and silica. The crystals were of various morphologies, including prisms, raphides, styloids, druses, crystal sand, spheres and clusters. Both calcium oxalate and calcium sulfate crystals were observed in almost all tissues, including mesophyll, parenchyma, sclerenchyma (fibre cells), pith, pith ray and cortex; calcium sulfate · magnesium oxalate crystals were only found in mesophyll and parenchyma cells in phyllodes.

Conclusions

The formation of most crystals was biologically induced, as confirmed by studying the crystals formed in the phyllodes from seedlings grown in a glasshouse. The crystals may have functions in removing excess calcium, magnesium and sulfur, protecting the plants against herbivory, and detoxifying aluminium and heavy metals.     

Medicago truncatula-derived calcium oxalate crystals have a negative impact on chewing insect performance via their physical properties

Plant structural traits often act as defenses against herbivorous insects, causing them to avoid feeding on a given plant or tissue. Mineral crystals of calcium oxalate in Medicago truncatula Gaertn. (Fabaceae) leaves have previously been shown to be effective deterrents of lepidopteran insect feeding. They are also inhibitors of conversion of plant material into insect body mass during or after consumption. Growth of beet armyworm, Spodoptera exigua Hübner (Lepidoptera: Noctuidae), larvae was correspondingly greater on calcium oxalate‐defective (cod) mutants of M. truncatula with lower levels of crystal accumulation. Data presented here show that insects feeding on M. truncatula leaves with calcium oxalate crystals experience greater negative effects on growth and mandible wear than those feeding on artificial diet amended with smaller amorphous crystals from commercial preparations. Commercial calcium oxalate can be added to insect artificial diet at levels up to 7.5‐fold higher than levels found in wild‐type M. truncatula leaves with minimal effect on insect growth or lepidopteran mandibles. These data suggest that negative impacts of calcium oxalate in the diet of larvae are due to physical factors, and not toxicity of the compound, as high levels of the commercial crystals are readily tolerated. In contrast to the dramatic physical effects that M. truncatula‐derived crystals have on insect mandibles, we could detect no damage to insect peritrophic gut membranes due to consumption of these crystals. Taken together, the data indicate that the size and shape of prismatic M. truncatula oxalate crystals are important factors in determining effects on insect growth. If manipulation of calcium oxalate is to be used in developing improved insect resistance in plants, then our findings suggest that controlling not only the overall amount, but also the size and shape of crystals, could be valuable traits in selecting desirable plant lines.     

Distribution of calcium oxalate crystals in floral organs of Araceae in relation to pollination strategy

? Premise of the study: Many flowers are pollinated by potentially hungry insects, yet flowers also contain gametes and embryos which must be protected from predation. Microscopic calcium oxalate crystals in plant tissues have been proposed to protect against herbivory. Aroids, which have an unusual diversity of such crystals, also exhibit diverse pollination strategies. Many species have pollinators that do not feed while visiting the flowers, while other species, especially those pollinated by beetles, offer sterile staminodia as food rewards. We examined flowers of 21 aroid species with various pollination strategies to test the hypothesis that crystals protect vital gametes and embryos while allowing consumption of food bribes. ? Methods: Aroid inflorescences collected from the field or from greenhouse material were sectioned, cleared, and examined by bright field and polarization microscopy. ? Key results: All species examined, regardless of pollination strategy, arrayed crystals around unshed pollen and ovules. Less vital tissues, such as odoriferous appendages, had few crystals. Staminodia offered as food to beetle pollinators, however, differed greatly between species in their crystal contents. Some had minimal crystals; some had crystals in patterns suggesting they limit beetle feeding; still others had abundant crystals in no obvious pattern. ? Conclusions: The results are consistent with crystals protecting against insect predation of gametes and embryos. However, the role of crystals in food-bribe staminodia is unclear. They may limit and direct feeding by beetles in some species, while in others they might have no protective role.     

Engineering calcium oxalate crystal formation in Arabidopsis

Many plants accumulate crystals of calcium oxalate. Just how these crystals form remains unknown. To gain insight into the mechanisms regulating calcium oxalate crystal formation, a crystal engineering approach was initiated utilizing the non-crystal-accumulating plant, Arabidopsis. The success of this approach hinged on the ability to transform Arabidopsis genetically into a calcium oxalate crystal-accumulating plant. To accomplish this transformation, two oxalic acid biosynthetic genes, obcA and obcB, from the oxalate-secreting phytopathogen, Burkholderia glumae were inserted into the Arabidopsis genome. The co-expression of these two bacterial genes in Arabidopsis conferred the ability not only to produce a measurable amount of oxalate but also to form crystals of calcium oxalate. Biochemical and cellular studies of crystal accumulation in Arabidopsis revealed features that are similar to those observed in the cells of crystal-forming plants. Thus, it appears that at least some of the basic components that comprise the calcium oxalate crystal formation machinery are conserved even in non-crystal-accumulating plants.     

Crystals in woody stems

Scanning electron microscopy has been used to examine the location and form of crystals in the stems of woody perennials. The results are discussed in relation to the effects of physical conditions and "impurities" upon crystal form. Infrared spectroscopy and X-ray diffraction have been used to establish the chemical identity of the crystals in a number of species which were chosen so as to provide a wide variation in form. Those from Populus deltoides and Polyalthia sp. consisted of calcium carbonate; those from ten other species consisted of hydrated calcium oxalate. The infrared spectra of the crystals from Dracaena fragrans, Terminalia balerica, Goniothalamus sp. and Combretum verticillatum were strongly indicative of calcium oxalate monohydrate, but in the other cases the degree of hydration of the oxalate was uncertain.     

Herbivory and Calcium Concentrations Affect Calcium Oxalate Crystal Formation in Leaves ofSida (Malvaceae)

Calcium oxalate crystals have potential roles in plants as partof a defence mechanism against herbivores and/or in accumulatingexcess calcium. To date, these potential roles have been studiedindependently. In this experimental study the effects of calciumlevels and herbivory on the production of calcium oxalate crystals(i.e. druse, spherical crystal aggregates) were examined inseedlings of Sida rhombifolia. Seedlings were subjected to threecalcium levels (low, normal or high) and an artificial herbivorytreatment. Calcium levels and herbivory both affected densityof crystals in leaves. Leaves from seedlings grown in low calciumhad a greater crystal density than those grown in high calcium.Leaves from seedlings subjected to herbivory had a greater crystaldensity than those from seedlings not subjected to herbivory.This study provides additional evidence that calcium oxalatecrystal production depends not only on calcium levels but canalso be influenced by external pressures such as herbivory.In addition to their physiological role in plants, these resultssuggest that calcium oxalate crystals can also act as a defencemechanism against herbivores. Copyright 2001 Annals of BotanyCompany Calcium concentrations, calcium oxalate crystals, herbivory, Malvaceae, Sida rhombifolia