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.     

Epidermal Structure and Stomatal Ontogeny in Some Polygonales and Centrospermae

The epidermal structure and ontogeny of stomata in 19 speciesof Centrospermae and two of Polygonales are described. The cellsof the epidermis are polygonal, isodiametric, or elongated invarious directions and arranged irregularly. The anticlinalepidermal walls are thick, sinuous, straight, or arched. Eleventypes of glandular and eglandular trichomes have been observed.Six types of stomata: anomocytic, paracytic, stomata with asingle subsidiary cell, diacytic, anisocytic, and transitionalbetween diacytic and paracytic, have been noticed in the speciesinvestigated. The ontogeny of anomocytic stomata is haplocheilicor perigenous, while that of the other five types is syndetocheilicor mesogenous. Abnormal stomata with a single guard cell, unequalguard cells, aborted guard cells, and arrested development arecommon. Groups of stomata are also frequent but contiguous stomataare rather rare.     

Structure,ontogeny and taxonomic significance of trichomes and stomata in some Capparidaceae

The present work embodies epidermal structure, structure and ontogeny of stomata in five genera embracing sixteen species of the Capparidaceae namely Cleome (8 species) Capparis (5 species), Cadaba (1 species), Crataeva (1 species) and Maerua (1 species). The epidermal cells are polygonal, isodiametric or elongated arranged irregularly, with evenly or unevenly thickened, sinuous, straight or arched anticlinal walls. Two main types of trichomes: glandular (four types) and eglandular (five types) are noticed. The stomatal types include cyclocytic, triacytic, staurocytic, tetracytic, anomocytic, anisocytic, paracytic and with a single subsidiary cell. The ontogeny of stomata with a single subsidiary cell is perigenous or mesoperigenous, of paracytic mesoperigenous or mesogenous, of anisocytic is mesoperigenous or mesogenous, while that of the other types is perigenous. Abnormalities observed are: single guard cell; aborted guard cells; complete or incomplete division of guard cells; contiguous stomata; giant stomata and cytoplasmic connections. The present observations do not support the separation of Cleomaceae from the Capparidaceae.     

Epidermal Structure and Ontogeny of Stomata in some Verbenaceae

The present paper deals with epidermal structure and developmentof stomata in 14 species of Verbenaceae. The epidermal cellsare either polygonal, isodia-metric, or elongated in variousdirections, and irregularly arranged. The anticlinal walls arethick, mostly sinuous, occasionally arched or straight. Thesurface of the cuticle shows parallel, rarely corrugated, striations.Some 12 types of eglandular and glandular trichomes, and foliarnectaries are noticed. The mature stomata are diacytic, anisocytic,paracytic, with a single subsidiary cell, anomocytic and perigenous.The development of anomocytic stomata is perigenous, while thatof others is mesogenous or syndetocheilic type. Abnormalitiesnoticed here include contiguous stomata, stomata with a singleguard cell, and aborted guard cells.     

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 genus Eugenia L. (Myrtaceae) in southern Africa: the structure and taxonomic value of stomata

The mature stomatal morphology of 11 southern African species of Eugenia , and also of E. uniflora L. (naturalized) and E. incerta Diimmer (a garden plant of unknown origin) has been studied by light and scanning electron microscopy.
Leaves of all the species are hypostomatic and water-stomata are occasionally present. Two different stomatal types, X and Y, differing mainly in their cuticular ornamentation, were found in the southern African species. The grouping of species based on stomatal characters supports a previous grouping of species on the basis of periderm and seed characters.
Conspicuous lipid bodies, usually present in the subsidiary cells, are limited to the southern African species. Although the subsidiary cells do not differ in shape and size from adjacent epidermal cells, their contents often have a higher tannin content than ordinary epidermal cells. The variable patterns of arrangement of the three to five subsidiary cells makes it difficult to identify the stomata with existing classifications based on mature topography. Thus, a new term 'anomostaurocytic' is proposed for the stomatal type found in the southern African species.
Stomata of E. uniflora and E. incerta are paracytic and anomocytic respectively, and the cuticular ornamentation of their stomata differs from those of the southern African species. The view that E. incerta is not closely related to the southern African species is supported by differences in its cuticular ornamentation, lack of lipid bodies in the subsidiary cells, anomocytic stomata and prominent T-pieces at the guard cell poles.     

Epidermal Structure and Stomatal Development in Some Malvaceae and Bombacaceae

Epidermal structure and development of stomata in 15 speciesof the Malvaceae and two of Bombacaceae are described. The cellsof the epidermis are polygonal, isodiametric, or elongated withthick, straight, arched, or sinuous anticlinal walls and containchloroplasts and abundant druses of calcium oxalate. Cuticularstriations radiating from the guard cells or hair bases arenoticed. Six types of glandular and non-glandular trichomesare seen. The mature stomata are anomocytic, anisocytic, andparacytic in the members investigated of both the families.The ontogeny of anisocytic and paracytic stomata is syndetochelicor mesogenous, while that of anomocytic is haplocheilic or perigenous.An abnormal stoma with a single guard cell is also observed.An increase in number of subsidiary cells in anisocytic stomatais due to the division of the subsidiary 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.     

Observations on the cotyledonary and hypocotyledonary stomata and trichomes in some Caesalpiniaceae with a note on their taxonomic

The structure and ontogeny of stomata on cotyledons and hypocotyls and the trichomes on hypocotyl are accounted for in eighteen species of Caesalpiniaceae. Trichomes are eglandular, bi- rarely tri-celled, smooth walled or walls wavy with cuticular striations or tubercles. Anomocytic, haplocytic, paracytic, diacytic, transitional, tetracytic, tricytic and cyclocytic stomata occur in different combinations even on the same surface of the cotyledon. In all, there are fourteen combinations. Inspite of the diversity, the most frequent type is anomocytic in most of the species and paracytic in some species of Cassia and Delonix (abaxial) but rarely it is haplocytic or anisocytic. In hypocotyls it is anomocytic. Ontogenetically anomocytic, tetracytic and cyclocytic stomata are perigenous, whereas other types are mesogenous or mesoperigenous. There is an increase in the number of subsidiary cells by their division or the neighbouring perigenes assuming their shapes. About eight such types are described. A pair of stomata has a common subsidiary cell. Twelve types of guard cell and stomatal approximation abnormalities are described. A range in the number, size and shape of the nuclei in guard cell are recorded. Megastomata (giant stomata) are observed in Parkinsonia and Tamarindus. The taxonomic significance of the stomata is also discussed.     

Morphogenetic effects of various growth substances on the cotyledonary stomata of brinjal and tomato

The effect of different growth substances on the development of normal and abnormal stomata are presented. Anomocytic, paracytic, anisocytic and stoma with a single subsidiary cell are observed. Abnormal developments like persistent stomatal cells, degeneration of guard cells, unusual thickening, unequal guard cells, single guard cells and size and shape of the pore are noticed in various growth substances. The growth substances also affect the stomatal frequency, stomatal index, epidermal frequency and size of guard and epidermal cells in both the plants. The highest meristematic activity is found in MOR 100 ppm in brinjal and in GA 25 ppm in tomato. The largest size of stomata is found in COL 25 ppm in brinjal and in MH 50 ppm in tomato. The same growth substance responds differently in the two plants.     

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)     

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.     

The stomatal apparatus of Lycopodium japonicum and its bearing on the stomata of the Devonian lycophyte Drepanophycus spinaeformis

The structure of the stomatal apparatus of the leaf of Lycopodium japonicum Thumb was studied using epidermal macerations, sections and scanning electron microscopy. The stomatal apparatus of L. japonicum consists of two large guard cells and pore, and is anomocytic. Based on light microscopy, the impression from epidermal macerations that there were two small guard cells surrounded by two, large, similarly shaped, subsidiary cells (paracytic) derives from a pronounced elliptical cuticular ledge on the surface of the guard cells surrounding a thickened circumporal area. A similar appearance is characteristic of cuticle preparations of the Devonian lycophyte Drepanophycus spinaeformis Göppert. We therefore conclude, as did W.H. Lang over 70 years ago, that the stomata of the early lycophyte were also anomocytic, as were those of a second species of Drepanophycus , D. qujingensis Li & Edwards.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 209–216.     

Stomatal development and patterning in Arabidopsis leaves

The functional unit for gas exchange between plants and the atmosphere is the stomatal complex, an epidermal structure composed of two guard cells, which delimit a stomatal pore, and their subsidiary cells. In the present work, we define the basic structural unit formed in Arabidopsis thaliana during leaf development, the anisocytic stomatal complex. We perform a cell lineage analysis by transposon excision founding that at least a small percentage of stomatal complexes are unequivocally non-clonal. We also describe the three-dimensional pattern of stomata in the Arabidopsis leaf. In the epidermal plane, subsidiary cells of most stomatal complexes contact the subsidiary cells of immediately adjacent complexes. This minimal distance between stomatal complexes allows each stoma to be circled by a full complement of subsidiary cells, with which guard cells can exchange water and ions in order to open or to close the pore. In the radial plane, stomata (and their precursors, the meristemoids) are located at the junctions of several mesophyll cells. This meristemoid patterning may be a consequence of signals that operate along the radial axis of the leaf, which establish meristemoid differentiation precisely at these places. Since stomatal development is basipetal, these radially propagated signals may be transmitted in the axial direction, thus guiding stomatal development through the basal end of the leaf.     

Structure,Ontogeny and Taxonomic Significance of Stomata in Some Cucurbitaceae

Stomatal structure, ontogeny in vegetative and floral organs of 9 genera and 12 species of Cucurbitaceae are described. The stomatal types conform to aperigenous, monoperigenous, diperigenous, hemipara-mesoperigenous and para-mesoperigenous types of Fryns-Claessens & Van Cotthem (1973). Stomatal abnormalities such as contiguous stomata, single guard cells with or without pore, one and a half stomata, degeneration of one or both the guard cells, cytoplasmic connections between guard cells of neighbouring stomata and a guard cell of a stoma and an adjacent epidermal cell, and division of guard cells are described. Stomata index, frequency of stomata, epidermal cells, size of guard and epidermal cells and organwise distribution of stomata are given. Stomatal studies does not support the view that the Cucurbitaceae are related to the Passifloraceae. The inclusion of 9 genera and 12 species studied in the tribe Cucumerineae is justified.     

Epidermal structure and development of stomata in vegetative and floral organs ofHybanthus enneaspermus (Linn.)F. Muell

The present paper deals with the epidermal structure and ontogeny of stomata in vegetative and floral organs ofHybanthus enneaspermus. The epidermal cells are either polygonal or elongated with straight, sinuous or arched thick anticlinal walls. The surface of the cuticle shows parallel striations radiating from the guard cells or hair bases. Unicellular and uniseriate bicellular trichomes with verrucose margins have been observed on all organs. The mature stomata are anisocytic, paracytic, anomocytic and transitional between anisocytic and paracytic. The ontogeny of anisocytic and paracytic stomata is syndetocheilic or mesogenous, anomocytic is haplocheilic or perigenous, while that of the transitional type is mesoperigenous. Four types of stomata have been observed on all the vegetative and floral organs and their ontogeny from organ to organ of this plant is constant. Stoma with a single guard cell is the result of disintegration of one of the guard cells before or after pore formation. Contiguous stomata are also occasionally noticed.     

The diversity of stomatal development in Orchidaceae subfamily Orchidoideae

The development of leaf stomata in species of Orchidoideae sensu Garay is agenous or hemimesogenous with a single mesogene cell. Both kinds of development occur in all 26 species studied and are found in a characteristic proportion which may differ markedly even between related species.
Leaf and stem stomata develop similarly but surrounding cells may divide obliquely in the latter, so that perigene cells are formed. At maturity, mesogene and perigene cells resemble other epidermal cells, the stomata being anomocytic.
In monocotyledons, the hemimesogenous development of stomata is previously known only from five orchid genera of the neottioid tribe Cranichideae sensu Dressler. This kind of stomatal development in monocotyledons is documented by micrographs for the first time. Mesogene and perigene cells are recorded for the first time in the Orchidoideae. The diversity of types of stomatal development in this group is emphasized.     

Structure and Development of Stomata in Some Labiatae

The structure and ontogeny of stomata have been studied in 33species of the Labiatae. The mature stomata are diacytic, transitionalbetween paracytic and diacytic, and anomocytic. The anomocyticstomata are haplocheilic or perigenous. The diacytic and thetransitional type of stomata are syndetocheilic or mesogenousas the two subsidiary cells and a pair of guard cells arisefrom the same meristemoid. The diacytic and the transitionalstomata are formed through three successive mitotic divisions.Abnormal stomata with single guard cells, arrested developments,and contiguous stomata have been observed. Contiguous stomataare formed either from two adjacently placed meristemoids orare the result of spatial readjustment during maturation ofthe leaf.     

The stomata of the cork-oak, Quercus suber. An ultrastructural approach

In the evergreen leaves of Quercus suber, stomata play a major role in adaptation to drought and temperature stress. The leaf is of zygostomic type and has about 430 stomata per square milimeter of abaxial leaf surface. The stomatal complex is of the anomocytic type. The guard cells protrude from the epidermal plane. The guard cell nucleus contains heterochromatin in small granules. The guard cell cytoplasm is characterised by a large number of well developed mitochondria, amyloplasts with stroma and grana, and a well developed cytoskeleton with a cortical array of microtubules oriented pa railed to the slit axis that persist even in mature cells. Guard cell walls are asymmetrically thickened and devoid of plasmodesmata. No area of cell walls was free of cuticle or covered by a thin cuticular layer and apparently no area of limited cuticular development provides evaporation when the stomata are closed.