Studies in Stomata of Chilli and Brinjal

Ontogeny and development of stomata in chilli (Capsicum annuumL.) and brinjal (Solanum melongena L ) are described Paracytic,anomocytic, and anisocyuc stomata are present Abnormalitiessuch as stoma with a single guard cell and degenerated stomatalcells are common. Degeneration of guard cell is traced in chilliJuxtaposed, superposed, and obliquely placed contiguous stomataare described. An incompletely differentiated menstemoid wasfound contiguous with a normal stoma Rarely a guard cell dividesto form a menstemoid. Cytoplasmic connections between the guardcells of adjacent or juxtaposed contiguous stomata are observedin chilli Extruded nucleoh in the guard cells of chilli arecommon Extruded nucleoh may enlarge The wounded epidermis ofchilli lose sinuosity of their walls and the degeneration ofguard cells is common.     

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.     

Some Observations on the Diversity of Stomata and Trichomes in Six Species of Dioscorea

Stomata and trichomes are described on the leaves of six speciesof Dioscorea. As many as six types were found in D. bulbiferaand D. oppositifolia, four types in D.hispida and D. wallichii,three types in D. belophylla, but only two types in D. alata.Although there is a diversity of stomata even on the same surface,the predominant type is anomocytic in all the species exceptD. bulbifera in which it is tricytic. Rarely a stoma is alsocyclocytic in D. bulbifera. An increase in the number of subsidiarycells in paracytic, tricytic, or diacytic stomata takes placeby the wall formation in them. Similarly a reduction in thenumber of subsidiary cells of a tetracytic stoma is the resultof lateral subsidiary cells assuming the form of epidermal cells.Abnormalities such as a stoma with one guard cell, degenerationof guard cells, and contiguous stomata are also met with. Theorganization of different types of stomata is studied in D.bulbifera and D. wallichii and shown to be perigenous. Capitateglandular hairs were seen on the leaves of D. belophylla, D.bulbifera, D. hispida, and D. wallichii but non-glandular, uniseriate,3-celled trichomes were observed only in D. hispida.     

Functional Stomata in a Variegated Leaf Chimera of Pelargonium zonale L. without Guard Cell Chloroplasts

Fluorescence microscopy indicated that chlorophyll was absentfrom epidermal and guard cells overlying all white areas andgreen areas (of certain leaves) in variegated leaves of Pelargoniumzonale, cv. Chelsea Gem. Stomata with chlorophyll-free guardcells, in general, responded normally to light and CO₂ as gaugedby direct measurements of stomatal aperture and by transpirationalwater loss studies, although stomata from white regions of variegatedleaves were more reluctant to open than stomata from green regionsof the leaves. Thus, functional stomata without guard cell chloroplastshave been discovered in another genus, namely Pelargonium, besidesthat originally discovered in Paphiopedilum. When stomata withchlorophyll-free guard cells opened, K⁺ accumulated in the guardcells. This indicates that chloroplasts are not essential forthe normal functioning of stomata and that the energy sourcefor driving stomatal movements can come from sources other thanphotophosphorylation. Key words: Guard cell chloroplasts, Leaf chimera, Pelargonium, Stomata     

Cellular Interactions in the Development of Stomatal Patterns in Vinca major L

The development of the leaf epidermis of Vinca major L. wasfollowed in situ by epi-illumination microscopy and evidencewas sought for cellular interactions. Stomata were often foundto be initiated in adjoining cells. The epidermal cells whichseparated such stomata when they had matured were formed fromthe same cells as the stomatal complexes themselves. The presenceof developing and mature stomata may influence only the orientationof divisions in neighbouring cells, and not the initiation andmaturation of stomata. There is great variability in the relativeorientation, timing and number of divisions which intervenebetween the first unequal division and the maturation of a stomaas well as the location of stomata relative to the spongy mesophylland minor veins. The results indicate that continuous short-rangeinteractions between the future guard cells and the adjoiningcells, rather than interactions between future stomata or afixed programme of development, are essential for the formationof the pattern of functional stomata in the mature leaf. Vinca major L., cell lineage, cellular interactions, development of stomata, epi-illumination microscopy, meristemoids, patterned differentiation, stomata     

Action of growth regulators on the cotyledonary stomata ofCucumis sativus L.: Structure and ontogeny

Anomocytic stomata and stomata with single subsidiary cells are commonly observed Sometimes a stoma appears anisocytic. Double cytoplasmic connections between nearby stomata and division of guard cells with persistent or degenerating nuclei are seen in GA. One or more divisions of guard cells, displaced guard cells and single guard cells with or without pore are noticed in SUC. Formation of single guard cells is a common feature in TIBA. Paracytic stomata, one and a half stomata and persistent stomatal initials are seen in SUL. COUM seems to be not inhibitory inCucumis sativus. In COL stomata with unequal guard cells, unequal stomatal cells with thickening in between but without intervening pore, stoma with double pores, persistent stomatal initials which may be solitary or in groups with varying shapes and with one or two nuclei of different shapes are noticed. The growth regulators affect the frequency of stomata, epidermal cells; stomatal index; size of guard and epidermal cells.     

The development of stomata and other epidermal cells on the rice leaves

In the leaves of rice (Oryza sativa), stomatal initials arose from two asymmetric cell divisions and a symmetric division. Guard mother cells (GMCs) and long cells in stomatal files (LCSs) were formed through the first asymmetric division of the precursor cell of GMCs. Subsidiary cells (SCs) were produced by the second asymmetric division of subsidiary mother cells or LCSs. Following SC formation, GMCs divided once symmetrically to generate guard cells and then differentiated terminally to form mature stomata. The developmental patterns of long cells, prickle hairs and short cells (phellem cells and silica cells) were also examined. Interestingly, we found that the different developmental stages of stomata and epidermal cells occurred in the similar location of immature leaves of the same phyllotaxis. In addition, two spacing patterns (“one stoma, one long cell” and “one short cell row”) probably exist in rice leaves.     

三种国产阴地蕨科植物叶的表皮构造

运用光学显微镜和扫描电镜对三种国产阴地蕨科植物：劲直蕨萁(Botrypus strictus)、小阴地蕨(Botrychium lunaria)和薄叶阴地蕨(Sceptridium daucifolium)〖WTBZ〗叶的成熟表皮构造进行了详细观察和研究。它们具有共同的特征：气孔散生,气孔类型无规则型,气孔长轴方向多与叶脉延伸方向一致。但也存在明显区别,特别是劲直蕨萁与另外二个种的区别更为明显：前者表皮细胞垂周壁直,相邻气孔不接触,保卫细胞平周壁具细条纹。但后二者之间亦有一定区别：薄叶阴地蕨的气孔为下生式,不下陷,而小阴地蕨的气孔为两面气孔型,气孔下陷。本文是国内首次对阴地蕨科叶的表皮构造进行研究,其研究结果表明,表皮构造在阴地蕨科植物的鉴定上具有重要意义,并且在研究阴地蕨科的分类以及起源和演化上也有一定参考价值。     

Structure of Polymorphic Stomata in Canella winterena (L.) Geartn. (Canellaceae)

The presence of 10 types of normal stomata along with 4 subtypes, one intermediate type between stephanocytic and hemiparacytic and some abnormal stomatal structures both in the vegetative and floral organs of Canella winterena (L.) Geartn. (Canellaceae) is reported here. The epidermal cells are polygonal, isodiametric in surface view with straight anticlinal walls. The stomata are amphibrachyparacytic, anomocytic, anisocytic, brachyparacytic, brachyparatetracytic, cyclocytic, hemiparacytic, laterocytic, holoparacytic, stephanocytic, one intermediate type between stephanocytic and hemiparacytic, etc. Abnormalities like contiguous stomata, different types of twin stomata and cytoplasmic bridge between adjacent stomata, stomata with single guard cell, single guard cell with pore juxtaposed with normal stoma are also found. The presence of giant stomata is a significant finding. (© 2014 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Structure and Development of Stomata in Some Leptosporangiate Ferns

The structure and development of stomata are described for 17species of leptosporangiate ferns. In these species the maturestomata are anomocytic, diacytic, Pyrrosia(applied), tetracytic,suspended, or floating type. Stomata are present on both surfacesof floating as well as sub merged leaves of Azolla and Marsilea.In a few species twin stomata, arrested development, and persistentstomatal initials are present Floating stomata result from disintegrationof the anticlinal suspending wall in Pleopeltis, and by detachmentand displacement in Azolla pinnata. The development of stomatais haplocheilic, syndetocheilic, or syndetohaplocheilic. InCyathea spinulosa the guard cell nuclei divide amitoticallyand the resulting two daughter nuclei occupy the opposite polesof the guard cells     

Structure and Ontogeny of Stomata in Some Polemoniales

The ontogeny and structure of stomata in 22 genera and 51 speciesof the Polemoniales are described. Five main types of stomatanoticed are: anisocytic, anomocytic, diacytic, paracytic, andstomata with a single subsidiary cell. Three modes of stomataldevelopment: syndetocheilic or mesogenous, haplocheilic or perigenous,and meso-perigenous or syndeto-haplocheilic are observed. Abnormalitiesseen are: stomata with single guard cells, arrested developmentand contiguous stomata variously oriented. Contiguous stomataresult from adjacently placed meristemoids or readjustment duringmaturation. Stomata with a single guard cell are formed as aresult of degeneration of one of the guard cells before or afterpore formation. The stomatal apparatus varies in different organsof a plant in form, number, orientation and arrangement of thesubsidiary and also the surrounding cells. Three lines leadingto Polemoniales, Boraginales, and Solanales are distinet.     

Arrangement of Stomata on the Leaves of Pelargonium zonale and Sedum stahlii

Stomata of Pelargonium zonale are orderly arranged on the surfaceof the lower epidermis. Inter-stomatal and Inter-trichome distancesas well as the number of sides of stomata, epidermal cells adjacentto stomata, and regular cells of the epidermis have characteristicprobability distributions. A model of epidermal developmentis constructed based upon the quantitative features of eachtype of cell. The primary restriction of the model is that stomainitials inhibit adjacent cells from becoming stoma initials. A second type of stomatal arrangement occurs in Sedum stahliiand an appropriate model for the ontogeny of epidermal cellsin this plant can also be constructed that has similar quantitativecharacteristics to those of real fields of cells. The applicationof modelling in plant morphology is discussed with respect tounderstanding ontogeny and phylogenic relationships.     

The Arabidopsis leucine‐rich repeat receptor‐like kinase MUSTACHES enforces stomatal bilateral symmetry in Arabidopsis

Stomata display a mirror‐like symmetry that is adaptive for shoot/atmosphere gas exchange. This symmetry includes the facing guard cells around a lens‐shaped and bilaterally symmetric pore, as well as radially arranged microtubule arrays that primarily originate at the pore and then grow outwards. Mutations in MUSTACHES (MUS), which encodes a leucine‐rich repeat receptor‐like kinase, disrupt this symmetry, resulting in defects ranging from skewed pores and abnormally focused and depolarized radial microtubule arrays, to paired guard cells that face away from each other, or a severe loss of stomatal shape. Translational MUSproMUS:tripleGFP fusions are expressed in cell plates in most cells types in roots and shoots, and cytokinesis and cell plates are mostly normal in mus mutants. However, in guard mother cells, which divide and then form stomata, MUS expression is notably absent from new cell plates, and instead is peripherally located. These results are consistent with a role for MUS in enforcing wall building and cytoskeletal polarity at the centre of the developing stoma via signalling from the vicinity of the guard cell membrane.     

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     

Structure and Development of Stomata in Vegetative and Floral Organs of Three Species of Kalanchoe

The structure and ontogeny of stomata in vegetative and floralorgans of three species of Kalanchoe is described. The matureanisocytic stomata are mono-cyclic or completely or incompletelyamphicyclic, rarely paracytic, transitional between paracyticand anisocytic and with a single subsidiary cell. The developmentof all the types is syndetocheilic or mesogenous from organto organ but the mature stomatal apparatus varies from organto organ as regards the number and arrangement of subsidiarycells. Abnormal stoma with a single guard cell and arresteddevelopment were observed on all organs. An abnormal stoma witha single guard cell develops directly from the meristemoid.     

Structure and Development of Stomata on the Vegetative and Floral Organs in Some Members of Caesalpiniaceae

The structure and development of stomata in 19 species of thefamily Caesalpiniaceae are described. The study is mostly confinedto the leaves, but observations have also been made on othervegetative and floral organs of some species Stomata may beparacytic, anisocytic, anomocytic, and with one subsidiary cell.Occasionally a stoma is diacytic, cyclocytic, or actinocytic.Different types occur individually or may be found side by sideeven on the same surface of an organ. The most prevalent typein all the genera is paracytic except in Caesalpima where itis anomocytic. The development of an anomocytic stoma is perigenous,but those with subsidiary cells are largely mesogenous; rarelyparacytic stomata are mesoperigenous. In spite of diversityof stomata, different types of stomata have similar patternsof development in different organs of the same plant. The presentinvestigation also indicates that the inconstancy of stomatain the family is due to (a) their diversity and (b) an increasein the number of subsidiary cells either by their division orby the neighbouring perigenes becoming subsidiary cell-like.     

叶子花表皮特征的研究及意义

对叶子花（Bougainvillea spectabilis）正常叶和变态叶上气孔密度、气孔指数和保卫细胞大小进行了研究。结果表明：正常叶上表皮的表皮细胞为多边形,垂周壁平直;下表皮的表皮细胞为不规则型,垂周壁浅波状;气孔类型为不规则型。变态叶上表皮没有发现气孔,变态叶下表皮的表皮细胞垂周壁则由浅波形逐渐变为深波形,气孔类型为不规则型和轮列型。随着变态叶的发育,变态叶下表皮的气孔密度降低,气孔指数升高;变态叶保卫细胞的长增大,宽减小。变态叶的平均气孔密度和平均气孔指数明显低于正常叶。正常叶和变态叶的保卫细胞均呈肾形。     

The Structure and Orientation of Guard Cells in Plants Showing Stomatal Responses to Changing Vapour Pressure Difference

Transmission electron micrographs revealed that a substantialpart of the guard cell wall of both Quercus robur L. and Populusnigra ‘italica’ L. was either free of cuticle orcovered with a greatly reduced cuticular layer. In Quercus thestructure of the guard cell was such that the area of limitedcuticular development would be exposed to the evaporating powerof the atmosphere even when the stomata were closed. Lanthanumstaining confirmed that this area might be an important siteof evaporation. A similar evaporation site was identified inthe guard cell wall of Pinus sylvestris L. Light micrographsrevealed that this area could also be exposed on the outsideof the leaf when the stomata were closed. It appears that guardcell orientation with respect to the epidermal plane dependsupon epidermal turgor. Changes in orientation of the guard cellcoupled with the exact location of the cuticle-free area inthe guard cell wall may explain the nature of the stomatal responseof individual species to changing VPD and the effect of othervariables, e.g. water deficit, on this response. Quercus robur L, oak, Populus nigra L, poplar, stomata, guard cells, cuticle, evaporation, vapour pressure difference     

合意非洲铁羽片表皮及气孔器的发育研究

用光学显微镜和扫描电镜对合意非洲铁羽片表皮及气孔器的发育过程进行观察研究。结果显示,(1)在羽片发育过程中表皮细胞变化不明显,主要是细胞伸长和壁增厚的过程;表皮的角质层连续出现。(2)气孔器发育可大致分3个阶段:保卫细胞母细胞阶段、幼保卫细胞阶段和成熟保卫细胞阶段;气孔器发育方式为单唇型或周源型;成熟气孔器属单环型。(3)在羽片发育过程中,气孔密度起初很低,后急剧升高并达到峰值,之后逐渐下降并趋于稳定。(4)有早熟气孔器。(5)羽片表皮发育成熟的方式是由基部向顶部逐步推进。     

An Unusual Feature of Stomatal Microanatomy in Certain Taxonomically- related Eucalyptus spp

The line of closure in the stoma in fully grown adult leavesof certain eucalypts is formed by special ridges of cuticlecalled ‘stomatal bars’ developed at the line ofclosure itself or from upgrowth of the cuticle of the lowersurface of the guard cells. Stomatal bars, previously discoveredin three members of the informal group ‘Bisectae’are shown to be restricted to certain species in that group.Possession of stomatal bars may affect stomatal performancebut does not appear to be a general adaptive response to thehabitats of the species which possess them. More probably, thepossession of stomatal bars is an inherited character with taxonomicvalue. The presence of stomatal bars in other genera is discussed. Eucalyptus spp, stomata, microanatomy, guard cells, stomatal bars