Studies on the structural and developmental variation and distribution of stomata in the Rubiaceae

The structure and ontogeny of the stomata has been studied in 26 species of Rubiaceae, in relation to their organographic distribution. The stomata are mostly paracytic on the leaves, but vary on other parts, particularly on the floral disk, where they are exclusively anomocytic. The foliar stomata are mesogenous and paracytic, but may be either dolabrate or trilabrate in origin. The disk stomata are perigenous and are probably derived from the mesogenous paracytic ones by the loss of divisive potential of their meristemoid due to anthogenetic factors. They are regarded to be hydathodal in function. If the dendroid habit is regarded as primitive, in the family, trilabrate stomata were quite possibly derived from dolabrate ones, which are mostly associated with the dendroid habit. Evidence from stomatal ontogeny is shown to be taxonomically important in the Rubiaceae.     

Structure,Delimitation,Nomenclature and Classification of Stomata

The paper reviews stomatal types observed in 500 species of angiosperms besides those described in the literature and deals with the problems of their structure, delimitation, nomenclature and classification. In view of the varied definitions available in the literature for subsidiaries, stomatal types and,the definition and delimitations being variously interpreted by different workers, a modified definition for the subsidiaries and stomata is presented. In accordance with the international code of nomenclature for plants, the names of the stomata widely in use are retained (rule of priority). They have been presently classified as pericytic, desmocytic, paracytic, diacytic, anisocytic, anisotricytic, isotricytic, tetracytic,staurocytic, anomocytic, cyclocytic and a good number of varieties under each type are presented. These stomatal types are recognised on the basis of their structure rather than its ontogenetic pathways.     

Structure, Delimitation, Nomenclature and Classification of Stomata

The paper reviews stomatal types observed in 500 species of angiosperms besides those described in the literature and deals with the problems of their structure, delimitation, nomenclature and classification. In view of the varied definitions available in the literature for subsidiaries, stomatal types and, the definition and delimitations being variously interpreted by different workers, a modified definition for the subsidiaries and stomata is presented. In accordance with the international code of nomenclature for plants, the names of the stomata widely in use are retained (rule of priority). They have been presently classified as pericytic, desmocytic, paracytic, diacytic, anisocytic, anisotricytic, isotricytic, tetracytic, staurocytic, anomocytic, cyclocytic and a good number of varieties under each type are presented. These stomatal types are recognised on the basis of their structure rather than its ontogenetic pathways.     

Stomatal patterning in Tradescantia: an evaluation of the cell lineage theory.

The cell lineage theory, which explains stomatal patterning in monocot leaves as a consequence of orderly divisions, was studied in Tradescantia. Data were collected to test the theory at three levels of organization: the individual stoma; stomata distributed in one dimension, in linear fashion along cell files; and stomata apportioned in two dimensions, across the length and breadth of the leaf. In an attempt to watch the patterning process through regeneration, stomata in all visible stages of development were laser ablated. The results showed that the formation of stomatal initials was highly regular, and measurements of stomatal frequency and spacing showed that pattern was determined near the basal meristem when the stomatal initials arose. Following the origin of initials, the pattern was not readjusted by division of epidermal cells. Stomatal initials were not committed when first present and a small percentage of them arrested. The arrested cells, unlike stomata, were consistently positioned in cell files midway between a developed pair of stomata. At the one-dimensional level of pattern, stomata in longitudinal files were separated by a variable number of epidermal cells and the frequency of these separations was not random. The sequential spacing of stomata also was not random, and stomata separated by single epidermal cells were grouped into more short and long series than expected by chance. The stomatal pattern across the width of the leaf resulted from cell files free of stomata which alternated with cell files containing stomata, but not with a recurring periodicity. Files lacking stomata were found only over longitudinal vascular bundles. Laser ablations of developing stomata did not disrupt the pattern in nearby cells or result in stomatal regeneration. We conclude that the cell lineage theory explains pattern as an individual stomatal initial arises from its immediate precursor and satisfactorily accounts for the minimum spacing of stomata in a cell file, i.e., stoma-epidermal cell-stoma. However, the theory does not explain the collective stomatal pattern along the cell files, at the one-dimensional level of patterning. Nor does the theory account for the for the two-dimensional distribution of stomata in which regions devoid of stomata alternate with regions enriched with stomata, but not in a highly regular nor haphazard manner. We suggest that the grouping of epidermal cells and stomata separated by single epidermal cells in cell files may result from cell lineages at a specific position in the cell cycle as they traverse the zone where stomatal initials form.(ABSTRACT TRUNCATED AT 400 WORDS)     

StomataStructure, Delimitation, Nomenclature and Classification of Stomata

The paper reviews stomatal types observed in 500 species of angiosperms besides thosedescribed in the literature and deals with the problems of their structure, delimitation, nomenclature andclassification. In view of the varied definitions available in the literature for subsidiaries, stomatal types and,the definition and delimitations being variously interpreted by different workers, a modified definition forthe subsidiaries and stomata is presented. In accordance with the international code of nomenclature forplants, the names of the stomata widely in use are retained (rule of priority). They have been presentlyclassified as pericytic, desmocytic, paracytic, diacytic, anisocytic, anisotricytic, isotricytic, tetracytic,staurocytic, anomocytic, cyclocytic and a good number of varieties under each type are presented. Thesestomatal types are recognised on the basis of their structure rather than its ontogenetic pathways.     

Stomatogenesis in the genus Hibiscus L. (Malvaceae)

In Hibiscus , stomata are anisocytic, anomocytic, paracytic and tetracytic, the first type being the most frequent and occurring in all plant parts in the ten species studied, whereas the others are scarce and have a limited organographic distribution. The stem, petiole, pedicel, staminal tube, ovary and style are stomatiferous; the leaf-blade, stipule, bracteole and sepals are amphistomatic and petals hypostomatic in the species investigated.
The stomatal types are often homoplastic, the anisocytic being either mesogenous trilabrate or mesoperigenous dolabrate, the anomocytic, mesoperigenous dolabrate or mesogenous trilabrate, and the tetracytic, mesoperigenous dolabrate or mesoperigenous trilabrate. But the typical paracytic stomata (with the subsidiaries completely enclosing the poles) are constantly mesogenous dolabrate and therefore probably indicate mesogenous dolabrate development. Although several patterns of stomatogenesis are encountered in any specific organ, only one of them is found to be dominant. A new subcategory of stomatal ontogeny, mesoperigenous trilabrate, is proposed in Hibiscus. No significant stomatal variation involving reduction in the divisive capacity of the meristemoid has been observed from the vegetative to floral parts; and stomata functioning as hydathodes have not been noticed in the latter, thus indicating that florogenic factors have no effect on the stomata.     

Epidermal Structure and Ontogeny of Stomata in Vegetative and Reproductive Organs of Ephedra and Gnetum

The epidermal structure and development of stomata in vegetativeand reproductive organs of Ephedra foliata and Gnetum ula isdescribed. The epidermal cells are polygonal, isodiametric,or elongated with thick or thin straight, arched, or slightlysinuous anticlinal walls. The cuticle is thin or thick. Papilla-likeunicellular outgrowths are present in Ephedra foliata. The maturestomata are orientated parallel to the longitudinal axes orirregularly. The mature stomata are anomocytic, paracytic, witha single subsidiary cell, cyclocytic, and actinocytic. Arresteddevelopment, contiguous stomata, and stomata with aborted guardcells have been observed. The ontogeny of stomata on differentorgans of these two plants is typically haplocheilic or perigenousbut the stomatal apparatus varies from organ to organ.     

Molecular phylogenetics, historical biogeography, and chromosome number evolution of Portulaca (Portulacaceae)

Portulaca is the only genus in Portulacaceae and has ca. 100 species distributed worldwide, mainly in the tropics and subtropics. Molecular data place the genus as one of the closest relatives of Cactaceae, but phylogenetic relationships within Portulaca are barely known. This study samples 59 species of Portulaca, 10 infraspecific taxa, and three cultivars, including multiple samples of widespread species. The sampled taxa represent all subgenera in the classifications of von Poellnitz (1934), Legrand (1958), and Geesink (1969) and come from around the world. Nuclear ITS and chloroplast ndhF, trnT-psbD intergenic spacer, and ndhA intron DNA sequences were analyzed using maximum likelihood and Bayesian methods to produce a hypothesis of relationships within Portulaca. Divergence times were estimated using Hawaiian endemics for calibration, and biogeographical patterns were examined using a Bayes-DIVA approach. In addition, the evolution of chromosome numbers in the genus was investigated using probabilistic models. The analyses strongly support the monophyly of Portulaca, with an age of the most recent common ancestor (MRCA) of 23 Myr. Within Portulaca are two major lineages: the OL clade (comprising opposite-leaved species) distributed in Africa, Asia, and Australia, and the AL clade (comprising alternate to subopposite-leaved species), which is more widespread and originated in the New World. Sedopsis, a genus sometimes recognized as distinct from Portulaca based on a long corolla tube, is nested within the OL clade and does not merit taxonomic recognition. Samples of Portulaca grandiflora, Portulaca halimoides, and Portulaca oleracea were found to be non-monophyletic. It is hypothesized that the ancestral distribution area of Portulaca included southern hemisphere continents and Asia. The OL clade remained restricted to the Old World (except Portulaca quadrifida, a pantropical weed), while the AL clade, with a South American origin, was able to disperse multiple times to other continents. The base chromosome number for Portulaca is inferred to be x=9, although the analysis was primarily based on the available data for the AL clade. A number of chromosome number change events (polyploidization, demi-polyploidization, gain, and loss) were shown to have occurred in the genus, especially within the Oleracea clade.     

COMPARATIVE ANATOMY OF THREE SPECIES OF THE APOPHYLLOUS GENUS GYMNOPHYTON

The anatomy of three species of Gymnophyton has been studied by light and electron microscopy. The species are essentially leafless and morphologically they are very much alike, but they differ anatomically and can be characterized just by their anatomical differences. SEM revealed great differences in stomatal structure, orientation, and dimensions. Micro-channels (ectodesmata) in the guard cell walls were disclosed in great numbers in G. isatidicarpum by using interference contrast microscopy, and these structures are thought to function as pathways for wax precursors. In older stems of this species the epidermis and cortical palisade tissue are isolated by a continuous periderm layer. Before dying, the palisade cells undergo alterations, and the stomata are permanently closed by cuticular plugs between the guard cells or by fusion of the swollen adaxial parts of the subsidiary cells. Similar permanent closure mechanisms are not found in G. polycephalum and G. robustum, which also deviate by having their stomatal openings orientated at a right angle to the axis of the stem and by the occurrence of collenchyma strands instead of fiber strands along the stem corner ribs. Gymnophyton polycephalum and G. robustum seem to be more closely related to each other than either is to G. isatidicarpum, but they differ markedly from one another in the size and structure of their stomatal complexes.     

Characteristics of Crassulacean Acid Metabolism in the Succulent C(4) Dicot, Portulaca oleracea L

Crassulacean acid metabolism (CAM) was investigated in leaves and stems of the succulent C₄ dicot Portulaca oleracea L. Diurnal acid fluctuations, CO₂ gas exchange, and leaf resistance were monitored under various photoperiod and watering regimes. No CAM activity was seen in well watered plants grown under 16-hour days. Under 8-hour days, however, well watered plants showed a CAM-like pattern of acid fluctuation with amplitudes of 102 and 90 microequivalents per gram fresh weight for leaves and stems, respectively. Similar patterns were also observed in detached leaves and defoliated stems. Leaf resistance values indicated that stomata were open during part of the dark period, but night acidification most likely resulted from refixation of respiratory CO₂. In water-stressed plants maximum acid accumulations were reduced under both long and short photoperiods. At night, these plants showed short periods of net CO₂ uptake and stomatal opening which continued all night long during preliminary studies under natural environmental conditions. Greatest acid fluctuations, in P. oleracea, with amplitudes of 128 microequivalents per gram fresh weight, were observed in water-stressed plants which had been rewatered, especially when grown under short days. No net CO₂ uptake took place, but stomata remained open throughout the night under these conditions. These results indicate that under certain conditions, such as water stress or short photoperiods, P. oleracea is capable of developing an acid metabolism with many similarities to CAM.     

外源硫对镉胁迫下马齿苋光合性状和矿质元素吸收的影响

从光合反应系统揭示外源硫(S)诱导马齿苋镉(Cd)耐受性的生理机制,为外源S缓解重金属毒害提供理论依据.采用营养液培养,研究外源S供体(NH4)2SO4对100 mg/L Cd胁迫下马齿苋叶片光合色素、光合特性、叶绿素荧光参数和矿质营养元素的影响.结果表明,Cd胁迫可显著降低马齿苋叶片中叶绿素a和叶绿素b含量;净光合速率、蒸腾速率、气孔导度均显著降低,而胞间二氧化碳浓度上升,表明非气孔因素是Cd胁迫诱导马齿苋光合抑制的主要因素;同时,PSⅡ实际光化学效率(ФPSII)、电子传递效率(J)、化学猝灭系数(qP)显著下降,而非化学猝灭系数(qN)显著上升,表明Cd胁迫影响马齿苋PSⅡ反应系统的正常运行.外施400 mg/L(NH4)2SO4显著提高马齿苋叶片叶绿素a含量、叶绿素b含量和叶绿素a/b比值,增强马齿苋叶片光合作用和PSⅡ原初光化学反应量子效率.对5种与光反应系统密切相关的矿质元素含量进行分析发现,Cd处理显著增加马齿苋叶片中的Ca和Fe含量,显著抑制马齿苋对Mg、Mn和Cu的吸收.Cd胁迫下马齿苋叶片的变黄与Mg、Mn的亏缺有关,而与Fe缺乏无关;添加外源S可显著提升马齿苋叶片中Ca、Mg、Fe、Cu和Mn含量,从而增强Cd胁迫下马齿苋叶片的PSII反应系统功能.     

中国石灰岩专性凤仙花叶表皮特征及其分类学意义

通过光学显微镜和扫描电镜技术对中国产的8种凤仙花科植物叶表皮微形态特征进行了实验研究,其中6种产于石灰岩地区,另外2种作为对照.结果表明,中国石灰岩地区的6种凤仙花的叶表皮微形态上、下表皮差异明显,上表皮细胞为不规则形和多边形,一般不具气孔器;下表皮细胞均为不规则形,均具气孔器,气孔器多为不等型.上、下表皮细胞形状、垂...     

The effect of exogenous abscisic acid on stomatal development, stomatal mechanics, and leaf gas exchange in Tradescantia virginiana

Gas exchange parameters and stomatal physical properties were measured in Tradescantia virginiana plants grown under well-watered conditions and treated daily with either distilled water (control) or 3.0 mM abscisic acid (ABA). Photosynthetic capacity (CO(2) assimilation rate for any given leaf intercellular CO(2) concentration [c(i)]) and relative stomatal sensitivity to leaf-to-air vapor-pressure difference were unaffected by the ABA treatment. However, at an ambient CO(2) concentration (c(a)) of 350 micromol mol(-1), ABA-treated plants operated with significantly lower c(i). ABA-treated plants had significantly smaller stomata and higher stomatal density in their lower epidermis. Stomatal aperture versus guard cell pressure (P(g)) characteristics measured with a cell pressure probe showed that although the form of the relationship was similar in control and ABA-treated plants, stomata of ABA-treated plants exhibited more complete closure at P(g) = 0 MPa and less than half the aperture of stomata in control plants at any given P(g). Scaling from stomatal aperture versus P(g) to stomatal conductance versus P(g) showed that plants grown under ABA treatment would have had significantly lower maximum stomatal conductance and would have operated with lower stomatal conductance for any given guard cell turgor. This is consistent with the observation of lower c(i)/c(a) in ABA-treated plants with a c(a) of 350 micromol mol(-1). It is proposed that the ABA-induced changes in stomatal mechanics and stomatal conductance versus P(g) characteristics constitute an improvement in water-use efficiency that may be invoked under prolonged drought conditions.     

马齿苋多糖提取物对翅果油树愈伤组织诱导的影响

采用传统热浸提工艺,从马齿苋植株中提取多糖.以B1培养基为基础,通过添加0.15 g/L的抗褐变剂Vc及不同浓度的马齿苋多糖,研究了其对翅果油树愈伤组织培养的抗褐变作用.结果显示:(1)马齿苋多糖对愈伤组织培养中的褐变有一定缓解作用.(2)经筛选,得到0.19 g/L为最适的多糖添加物浓度,子叶愈伤组织产生生物量最多,抗褐变效果与Vc的作用相似,愈伤组织生命周期也明显缩短.     

The bHLH protein, MUTE, controls differentiation of stomata and the hydathode pore in Arabidopsis

Stomata are turgor-driven epidermal valves on the surface of plants that allow for efficient gas and water exchange between the plant and its environment. The Arabidopsis thaliana basic helix-loop-helix (bHLH) protein, MUTE, is a master regulator of stomatal differentiation where it is required for progression through the stomatal lineage and the differentiation of stomata. The genetic control of stomatal spacing across the epidermal surface is variable in different organs. For instance, a distinct suite of genes from those in leaves regulates stomatal patterning in hypocotyls. Here we report that regardless of organ type, MUTE controls downstream events directing stomatal differentiation, specifically the transition from meristemoid to guard mother cell. Ectopic MUTE expression is sufficient to over-ride cell fate specification in cell types that do not normally differentiate stomata. Furthermore, MUTE is required for the production of the structure evolutionarily related to stomata, the hydathode pore. Consistently, MUTE displays expression at the tip of cotyledons and leaves, thus co-localizing with the auxin maxima. However, MUTE itself was not regulated by the auxin, and the absence of hydathode pores in mute did not affect the auxin maxima. Surprisingly, our analysis revealed that the requirement for MUTE could be partially circumvented under conditions of compromised inhibitory signaling.     

Stomatal patterning in angiosperms

My thesis is that understanding stomatal patterning requires a holistic perspective. Since stomata are structures critical to the survival of terrestrial plants, they need to be viewed in relation to their function and their interface with other structural components. With this outlook, I begin by discussing pattern types, means of measuring them, advantages of each type of measurement, and then present patterning from evolutionary, physiological, ecological, and organ views. I suggest areas where I believe profitable studies might enable us to better understand stomatal patterning. The final sections of the paper review stomatal patterning on angiosperm leaves and present a theory of patterning. With the abundance of molecular information, and coming genomic sequences and new tools, an opportunity exists to dissect the process of how cells are selected to become different from their neighbors and assume a fate critical to plant survival. Understanding this biological process at the molecular level requires comprehending the broad base on which stomatal patterning rests.     

热胁迫下不结球白菜和甘蓝叶片组织结构的变化

利用显微及扫描电镜方法,研究不结球白菜及甘蓝的耐热品种和感热品种种在高温（３８－３９℃）胁迫下,叶片表皮和组织结构的差异。结果如下：耐热品种叶片下表皮比感热品种气孔频度高,气孔体积小且开张度小。耐热品种部分气孔呈关闭状态,而感热品种气孔基本上呈开放状态。显微结构观察到耐热品种叶肉组织细胞排列紧密,叶脉的维管束发达,特别是木质部的导管数量多且孔径大。而感热品种相比于耐热品种叶肉细胞排列疏松,维管束相     

马齿苋多糖的研究

用苯酚硫酸法对马齿苋多糖含量进行测定,设计正交实验确定马齿苋多糖提取的最佳工艺,马齿苋多糖的提取率高达9.23%。将其多糖分离纯化,进行理化性质试验测试,利用纸层析和气相色谱对马齿苋多糖中的单糖组成做了分析,马齿苋多糖中的单糖组成有葡萄糖、半乳糖、甘露糖、果糖、木糖、阿拉伯糖。     

Stomata and Structure of Tetraploid Apple Leaves cultured in Vitro

Leaves of anther-derived tetraploid apple (Malus pumila Mill.)shoots were examined by low-temperature scanning electron microscopy(LT-SEM). Leaves were serrate and wide with an undulating adaxialsurface due to convex epidermal cells, apparently without crystallineepicuticular wax. Stomata were absent from the adaxial surface,except for the marginal teeth which exhibited 40-60 stomataper leaf; they probably originated from residual mitotic activity.One third of abaxial stomata was occluded by the residual cuticleof the mother guard cell across the stomatal pore which rupturedwhen the stomata became functional. The stomatal index was 7·2(± 1·6) with 60-75 stomata mm^-2, i.e. abaxialstomata of tetraploid leaves expanded in vitro were less frequentthan those in triploid leaves either cultured in vitro (475-575stomata mm^-2) or grown on the tree (320-390 stomata mm^-2) wherethe stomatal index was 21 (± 4). Freeze-fracture transsectionsshowed that the tetraploid in vitro leaves were composed ofa layer of adaxial epidermal cells, followed by a single layerof palisade cells and four to five layers of spongy mesophyllcells and the abaxial layer of epidermal cells, in contrastto juvenile seedling-grown apple leaves in which the two layersof palisade cells comprised the majority (52-60%) of the leafvolume. The same morphological features, such as wide and lesspointed leaves, reduced stomatal density and stomatal index,and increased stomatal size that were previously reported fortree-grown tetraploid leaves were also expressed in vitro. Thus,causes of the stomatal deformation in tissue-cultured Rosaceaeare interpreted to be in part genetic and not purely environmental.Copyright1994, 1999 Academic Press Malus pumila Mill., apple, biotechnology, breeding, cryo-preservation, CO₂, juvenile, low temperature-scanning electron microscopy (LT-SEM), micropropagation, ploidy, stomata, tissue-culture, transpiration