Stomatal sensing of the environment

The effects of environmental factors on stomatal behaviour are reviewed and the questions of whether photosynthesis and transpiration eontrol stomata or whether stomata themselves control the rates of these processes is addressed. Light affects stomata directly and indirectly. Light can act directly as an energy source resulting in ATP formation within guard cells via photophosphorylation, or as a stimulus as in the case of the blue light effects which cause guard cell H⁺ extrusion. Light also acts indirectly on stomata by affecting photosynthesis which influences the intercellular leaf CO₂ concentration ( C _i). Carbon dioxide concentrations in contact with the plasma membrane of the guard cell or within the guard cell acts directly on cell processes responsible for stomatal movements. The mechanism by which CO₂ exerts its effect is not fully understood but, at least in part, it is concerned with changing the properties of guard cell plasma membranes which influence ion transport processes. The C _i may remain fairly constant for much of the day for many species which is the result of parallel responses of stomata and photosynthesis to light. Leaf water potential also influences stomatal behaviour. Since leaf water potential is a resultant of water uptake and storage by the plant and transpirational water loss, any factor which affects these processes, such as soil water availability, temperature, atmospheric humidity and air movement, may indirectly affect stomata. Some of these factors, such as temperature and possibly humidity, may affect stomata directly. These direct and indirect effects of environmental factors interact to give a net opening response upon which is superimposed a direct effect of stomatal circadian rhythmic activity.     

Structure and development of stomata on the primary root of Ceratonia siliqua L

Stomata of various sizes are produced on the primary root of Ceratonia siliqua L. Most are generated during embryogenesis, prior to seed desiccation. They can be detected on the dry embryo in a wide zone just above the root tip. Initially, large stomata are formed. These have the ability to induce divisions of their neighbouring cells, creating particular cell patterns around them. Later, small perigenous stomata are generated. As the root grows following seed germination, the stomatal zone overlaps with that of the root hairs. Although root stomata of C. siliqua undergo a structural differentiation that seems almost identical to that of the elliptical stomata formed on leaves, they are unable to move and remain permanently open. Polarizing microscopy of fully differentiated stomata and young stomata at the stage of stomatal pore formation revealed deposition of radial cellulose microfibril systems on their periclinal walls. However, these systems were less developed than those on leaf stomata, a feature that might be responsible for their inactivity. Besides, plastids of the root guard cells (GCs) do not differentiate into chloroplasts but function solely as amyloplasts. Root stomata have a short life span. During rapid and intense root growth, GCs cannot keep pace with the elongation of their neighbouring rhizodermal cells. They therefore split in their mid-region, transversely to the stoma axis. The two parts of the transversely torn stoma are dragged apart and a large opening is formed on the root surface, just above the substomatal cavity. The root stomata, together with these openings, may facilitate increased gaseous exchange during respiration and/or an increased transfer of some nutrients and water in the rapidly growing primary root.     

A scanning electron microscope study of normal and vitrified leaves from Datura insignis plantlets cultured in vitro

The surface anatomy of normal and vitreous leaves of plantlets obtained from Datura insignis Barb Rodr nodal segment cultures was compared using scanning electron microscopy. Normal and vitrified leaves are similar in several ways. They are both amphistomatic, and have similar distributions of glandular and non-glandular trichomes. Stomata have similar length, diameter and distribution in normal and vitreous plants. Immature stomata, which have closed pores, and plugged stomata, which contain an amorphous material between their guard cells, occur in both normal and vitrified leaves. Normal and vitreous leaves differ in the frequency of normal and abnormal stomata. Normal stomata have kidney-shaped guard cells and resemble closely those found in field-grown plants, whereas abnormal stomata have deformed guard cells. Normal stomata represent approximately 80% of the total number of stomata in normal leaves, but only 7% of the total number of stomata in vitreous leaves. Abnormal stomata represent 90% of the total number in vitreous leaves. The deformation of guard cells could possibly be a mechanical impediment to stomatal function.     

Zooming into sub-organellar localization of reactive oxygen species in guard cell chloroplasts during abscisic acid and methyl jasmonate treatments

Regulation of stomata movements is crucial for plants ability to cope with their changing environment. Guard cell’s (GC) water potential directs water flux inside/outside this cell, which eventually is causing the stoma to open or close, respectively. Some of the osmolytes which accumulates in the GC cytoplasm and are known to play a role in stomata opening are sugars, arising from chloroplast starch degradation. During stomata closure, the accumulated osmolytes are removed from the GC cytoplasm. Surprisingly little is known about prevention of starch degradation and forming additional sugars which may interfere with osmotic changes that are necessary for correct closure of stomata.   One of the early events leading to stomata closure is production of reactive oxygen species (ROS) in various sub-cellular sites and organelles of the stoma. Here we report that ROS production during abscisic acid (ABA) and methyl jasmonate (MJ) stimuli in Arabidopsis GC chloroplasts were more than tripled. Moreover, ROS were detected on the sub-organelle level in compartments that are typically occupied by starch grains. This observation leads us to suspect that ROS function in that particular location is necessary for stomata closure. We therefore hypothesize that these ROS are involved in redox control that lead to the inactivation of starch degradation that takes place in these compartments, thus contributing to the stoma closure in an additional way.     

Osservazioni Sulla Maggiorana (Origanum Majorana L.) in Veste di Pianta Coltivata

Abstract

Stomatal features and ontogeny of stomata of 11 ornamental taxa of monocotyledonous families with Agavaceae (1 species), Amaryllidaceae (1 species), Araceae (3 species), Cannaceae (1 species), Commelinaceae (3 species), Liliaceae (1 species), and Musaceae (1 species) have been studied. Features like stomatal area, leaf area occupied by stomata and per cent leaf area occupied by stomata are reported for these taxa for the first time.     

Ecology and Morphology of Blossfeldia liliputana (Cactaceae): a Poikilohydric and almost Astomate Succulent*

Plants of the monotypic Blossfeldia liliputana have the smallest bodies of all Cactaceae. The button-like plants with a diameter of usually some 10 mm occur in rock crevices in arid regions between S Bolivia and N Argentina. Based on observations and experiments in the field and in cultivation, morphology, anatomy, reproductive biology, certain aspects of ecophysiology, and behaviour under water stress are described. The very small flowers are autogamous; the arillate hairy seeds are unique within the family and represent a particular adaptation to ant dispersal. These CAM-plants virtually lack stomata: 0.6 stomata/mm² represents the lowest number in terrestrial autotrophic vascular plants. However, all other xeromorphic features characteristic for globular cacti are absent (e.g. no thickened cuticle, no thickended outer cell walls, no thickened hypodermal layers). These features allow a high degree of desiccation: under water stress the plants lose up to 80% of their weight within one year and can withstand an additional drought of at least another year. Thus Blossfeldia is poikilohydric like many lichens and mosses and represents the unique life form of a succulent resurrection plant.     

田间小麦叶片气孔对环境因子响应的模拟及叶片水分平衡的计算

经实测与数据分析得出田间供试小麦叶片气孔对光强、叶温、叶片水势分别作一定形式的响应,据此估算了冠层总气孔导度,进而建立了预测小麦群体蒸腾速率、冠层总气孔导度、叶温、叶片水势等的水分平衡模型。实验表明模型具有较好的预测性。供试小麦叶片一天中可溶性糖含量变化对叶片水势的影响不大。     

Cadmium effects on leaf transpiration of sugar beet (Beta vulgaris)

Seedlings of sugar beet ( Beta vulgaris L. cv Monohill) were cultivated for 4 weeks in nutrient solution containing different concentrations of CdCl₂ (0 to 10 μ M ). The effects of Cd on appearance and function of stomata and leaf cuticle were investigated by water loss measurements and microscopy. The leaf transpiration rate increased with increasing Cd concentrations while the sum total of stomatal aperture area per unit leaf area decreased. Already at low Cd levels. an increase of defective and undeveloped stomata was found in Cd treated plants. These stomata are closed or have small apertures and probably lack a functional closing mechanism. The number of intact stomata per unit leaf area was lower in leaves of Cd treated plants than in controls, and Cd induced closure of intact stomata. The total number of stomata per leaf area slightly increases with increasing Cd concentration. as does the percentage of small stomata. Furthermore. specific leaf area increased, while the density of leaf structure was decreased by Cd. From this observation we conclude that the increase in transpiration rate caused by Cd is primarily due to effects on the permeability of the leaf cuticle to water.     

The adaptive significance of amphistomatic leaves

Abstract A clear correlation between the presence of stomata on both surfaces and factors such as habitat, growth form, and physiology has yet to emerge in the literature. However, certain loose trends with these factors are evident, and these are reviewed along with evidence for hypostomaty as the primitive form. It is proposed that the effect of developing stomata on the upper surface as well as the lower is to increase maximum leaf conductance to CO₂. Plants with a high photosynthetic capacity, living in full-sun environments, and experiencing rapidly fluctuating or continuously available soil water (as opposed to seasonal or long-term soil water depletion), are identified as deriving an adaptive advantage from a high maximum leaf conductance. The correlation between groups of plants fitting the above conditions and those noted to be amphistomatic is remarkable. Plants not fitting the conditions are found to be largely hypostomatic.     

Variation in the structure and development of foliar stomata in the Euphorbiaceae

The structure and ontogeny of foliar stomata were studied in 50 species of 28 genera belonging to 17 tribes of the family Euphorbiaceae. The epidermal cells are either polygonal, trapezoidal, or variously elongated in different directions and diffusely arranged. The epidermal anticlinal walls are either straight, arched or sinuous. The architecture of cuticular striations varies with species. The mature stomata are paracytic (most common), anisocytic, anomocytic and diacytic. Occasionally a stoma may be tetracytic, cyclocytic or with a single subsidiary cell. The ontogeny of paracytic stomata is mesogenous dolabrate or trilabrate, mesoperigenous dolabrate; that of diacytic stomata is mesogenous dolabrate, whereas that of anisocytic stomata is mesogenous trilabrate; rarely an anisocytic stoma may be mesoperigenous. Hemiparacytic stomata are mesoperigenous unilabrate; tetracytic stomata are mesoperigenous dolabrate and anomocytic stomata perigenous. Abnormalities encountered include four types of contiguous stomata, stomata with a single or both guard cells aborted and persistent stomatal initials. Cytoplasmic connections between the guard cells of two adjacent stomata or the guard cell of a stoma and an adjacent epidermal/subsidiary cell, or both types occurring in a species, were noticed. The stomatal development, distribution, diversity and basic stomatal type with reference to systematics are discussed.     

骆驼蓬营养器官的旱生结构

 骆驼蓬是生长在荒漠或盐碱地区的旱生植物,具有显著的旱生结构特征。主根粗壮,其中柱周围产生2或3轮呈同心环状排列的异常维管束。这种异常结构对旱生植物具有重要的生态学意义。茎肉质,其皮层、韧皮部,尤其是髓内具有发达的贮水薄壁组织细胞。叶片肉质,光滑无毛。其表皮细胞的外切向壁有较厚的角质层,表面具皱纹状突起。气孔器与表皮细胞平齐,面积较大,而密度较小。栅栏组织发达,为环栅型,包围着发达的贮水组织。叶脉维管束不发达。其叶表面积与体积之比远比中生植物小。根据以上营养器官形态结构特征,可以认为骆驼蓬是典型的多浆汁旱生植物。     

水杨酸对黄瓜子叶表皮气孔开度的调节作用

以黄瓜品种中农203(Cucumis sativus L.cv.Zhongnong 203)幼苗为试材,采用SA溶液根部施用和子叶表皮浸泡两种方式,显微观测了不同外源水杨酸(Salicylic acid,SA)溶液处理对其子叶表皮气孔开度的影响,以探讨SA与气孔运动的关系.结果表明:SA子叶表皮浸泡或根部施用后,气孔运动的趋势是随着SA浓度增加而孔径逐渐变小,且SA磷酸缓冲液的作用效果与SA水溶液相似.随着处理时间的延长,气孔开度逐渐变小,且气孔开度与SA处理时间达极显著(r=-0.962**)或显著(r=-0.914*)负相关.溶液低pH值,增强了SA对气孔开度的抑制作用,且SA浓度越高作用越明显;0.1 mmol/L SA处理后,pH为8、7、6溶液的气孔开度抑制率分别为90.2%、93.8%和96.3%,即SA溶液对气孔开度的抑制率随着溶液pH降低而升高.可见,外源SA能够促进气孔关闭,其作用随着SA浓度升高、处理时间延长和溶液pH值降低而增强,相对于磷酸缓冲液,以蒸馏水作为溶剂的SA溶液促进气孔关闭的作用更大.     

Imbibition of Swietenia macrophylla (Meliaceae) seeds: the role of stomata

BACKGROUND AND AIMS: The occurrence of stomata in seed coats is uncommon and there is limited information about their function(s). The aim of this study was to verify the distribution of stomata in seed coats of Swietenia macrophylla and to relate it to the imbibition process and aspects of the structure of the outer integument layers. METHODS: For the structural and ultrastructural studies, the seeds were processed using the usual techniques and studied under light and scanning electron microscopes. Histochemical tests were employed to identify the cell wall composition in the different seed coat portions. To assess the role of the stomata in the imbibition, non-impervious seeds were compared with partially impervious ones, in which only the embryo, median or hilar regions were left free. Further, the apoplastic pathway marker was employed to confirm the role of the stomata as sites of water passage during imbibition. KEY RESULTS: A positive relationship was observed between seed coat thickness and stomata density. The stomata were devoid of movement, with a large pore. They occurred in large numbers in the embryo region and extended with lower frequency towards the wing. Imbibition rates were related to stomata density, suggesting that the stomata act as preferential sites for water entry in the S. macrophylla seeds. CONCLUSIONS: At maturity, the stomata in the seed coat play a significant role in seed imbibition. The data may also infer that these permanently opened stomata have an important role in gas exchange during seed development, aiding embryo respiration.     

European and Mediterranean oaks (Quercus L.; Fagaceae): SEM characterization of the micromorphology of the abaxial leaf surface

This paper examines micromorphological characteristics (trichomes, waxes, stomata) of the abaxial leaf surface of European and Mediterranean oaks. Observations were performed In-scanning electron microscopy, and the purpose of the investigation was to ascertain whether it is possible to distinguish the different species taxonomically using these features. Several species, as well as some subspecific entities and taxa considered doubtful, were taken into consideration. Subgenera appear to be well-defined: e.g. the subgenus Quercus is characterized by waxes arranged in vertical scales, the subgenera Cenis and Sclerophyllodris by smooth waxes. They differ because the former has markedly elliptical stomata, while the latter two have roundish stomata. The different species are also fairly easy to distinguish, and this fact confirms the taxonomical validity of the parameters we have taken into consideration. In some cases subspecific entities and doublful species do not diflcr at all from the related species, but in others they present such marked variations that it is difficult even to classify them with certainty. The most critical taxonomic groups (i.e. those which include a number of specific and subspecific entities often too difficult to distinguish from each other) are the series Quercus robur – Quercus petraea,– Quercus pubescens and the Quercus faginea,– Quercus Insitanica Quercus canariensis group.     

Stomatal responses to changes in humidity in plants growing in the desert

Summary The stomata of plants growing in the Negev Desert, namely the stomata of the mesomorphic leaves of Prunus armeniaca, the xeromorphic stems of Hammada scoparia, and the succulent leaves of Zygophyllum dumosum, respond to changes in air humidity. Under dry air conditions diffusion resistance increases. Under moist air conditions diffusion resistance decreases. When the stomata close at low air humidity the water content of the apricot leaves increases. The stomata open at high air humidity in spite of a decrease in leaf water content. This excludes a reaction via the water potential in the leaf tissue and proves that the stomatal aperture has a direct response to the evaporative conditions in the atmosphere. In all species the response to air humidity is maintained over a period of many hours also when the soil is considerably dry. The response is higher in plants with poor water supply then in well watered plants. Thus for field conditions and for morphologically different types of photosynthesizing organs the results confirm former experiments carried out with isolated epidermal strips.     

The guard-cell apoplast as a site of abscisic acid accumulation in Vicia faba L.

Abscisic acid (ABA) integrates the water status of a plant and causes stomatal closure. Physiological mechanisms remain poorly understood, however, because guard cells flanking stomata are small and contain only attomol quantities of ABA. Here, pooled extracts of dissected guard cells of Vicia faba L. were immunoassayed for ABA at sub‐fmol sensitivity. A pulse of water stress was imposed by submerging the roots in a solution of PEG. The water potentials of root and leaf declined during 20 min of water stress but recovered after stress relief. During stress, the ABA concentration in the root apoplast increased, but that in the leaf apoplast remained low. The ABA concentration in the guard‐cell apoplast increased during stress, providing evidence for intra‐leaf ABA redistribution and leaf apoplastic heterogeneity. Subsequently, the ABA concentration of the leaf apoplast increased, consistent with ABA import via the xylem. Throughout, the ABA contents of the guard‐cell apoplast, but not the guard‐cell symplast, were convincingly correlated with stomatal aperture size, identifying an external locus for ABA perception under these conditions. Apparently, ABA accumulates in the guard‐cell apoplast by evaporation from the guard‐cell wall, so the ABA signal in the xylem is amplified maximally at high transpiration rates. Thus, stomata will display apparently higher sensitivity to leaf apoplastic ABA if stomata are widely open in a relatively dry atmosphere.     

白刺叶片气孔特征对人工模拟降雨的响应

以荒漠生态系统典型植物白刺(Nitraria tangutorum Bobr)叶片为研究对象,利用数码图像显微镜处理系统,研究了不同人工模拟增雨处理下的白刺叶片气孔密度及形态特征的变化情况。结果表明,荒漠植物固有特征决定了白刺叶片下表皮气孔密度大于上表皮,上表皮、下表皮气孔密度对增雨响应差异不显著(P0.05)。增雨处理上表皮、下表皮气孔密度与对照差异显著(P0.05)。相同增雨季节,50%处理下叶片气孔密度高于100%处理;不同增雨季节,气孔密度对生长季后期增雨响应更明显。白刺叶表皮气孔分布遵循"一细胞间隔(one cell spacing rule)"法则。增雨后叶片上表皮和下表皮气孔长度、宽度均有不同程度的增加,气孔形态特征对100%处理的响应较50%处理更为明显,且生长季后期增雨对叶片气孔形态特征的影响更大。     

Comparative Studies on Water Relations and Xeromorphic Structures of Some Plant Species in the Middle Part of the Desert Zone in China

The mean value of leaf water potential (LWP) of the mesophytes, the less-succulent xerophytes and the succulent xerephytes was –13.5, –18.09 and –26.09 bars respectively. The bound water content (BWC), and the ratio of bound water content to free water content (B/E ratio) for the succulent xerophytes (174.0% and 4.8) were higher than those for the mesophytes (52.5% and 0.37) and the less-succulent xerophytes (76.8% and 1.0). The transpiration rate of the mesophytes, less-succulent xerophytes and succulent xerophytes was 1146.5, 866.3 and 422.8 mg/g. fw. h respectively. The water holding capacity (WHC) of the succulent xerophytes was the strongest among those of the three types, while that of the mesophytes was the weakest. The xerophytes, both the succulent and the less-succulent, have the following xeromorphic structures: thick cuticle, sundan stomata, dense epidermal hairs, well-developed palisade tissue and degenerated spongy tissue, the high ratio of palisade tissue to spongy tissue, and equilateral leaves. Besides, succulent xerophytes possess other xeromorphic structures: thick leaves, well-developed water storing tissue, high ratio of water tissue to leaf thickness, and the succulent leaves. The mesophytes exhibit the following mesophilous structures, thin cuticle, levelling or arching up stomata, low ratio of palisade tissue to spongy tissue, and back-belly leaves.