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 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.     

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.     

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.     

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.     

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.     

Trichomes and stomata,and their taxonomic significance in theUrticales

Structure and ontogeny of stomata and trichomes have been studied in 23 species and 3 varieties of theUrticales. Stomata are anomocytic, more rarely paracytic; anisocytic and sometimes helicocytic and transitorial types are found inUrticaceae andDorstenia, rarely inArtocarpus. The ontogeny of anomocytic and actinocytic stomata is perigenous, of paracytic either mesogenous or perigenous, of anisocytic either mesogenous or mesoperigenous, and of helicocytic and transitional types mesogenous. Among trichomes eglandular unicellular (wide spread), bicellular or uniseriate filiform (Cannabis); glandular capitate with uni- or bicellular (Moraceae, Ulmaceae, Cannabaceae), uniseriate filiform (Ulmaceae) or multiseriate stalk (Cannabis); sunken glands (Artocarpus); uniseriate glandular with uniseriate stalk (Celtis), and stinging emergences (Urticaceae) have been observed. It is concluded that theUrticales represent a natural order with four families:Ulmaceae, Moraceae, Urticaceae andCannabaceae which are distinct but interrelated with each other.     

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.     

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.     

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.     

Observations on the structure and ontogeny of stomata and trichomes on developing and mature pericarps ofCassia occidentalis L

Paracytic, anisocytic, anomocytic, transitional forms, tetracytic, cyclocytic stomata and partly and completely amphicyclic forms are found, often on the same surface, in nine combinations. The most frequent type is paracytic. A few morphological variations in the basic types and eight types of abnormalities in stomata are recorded. The stomatal ontogeny may be mesogenous, mesoperigenous or perigenous. Trichomes are multicellular glandular club-shaped and unicellular eglandular. The taxonomic significance of stomata is indicated.     

Observations on the effects of morphactin EMD 7301 and EMD 7311 on the number,size, morphology and ontogeny of cotyledonary stomata ofAbelmoschus esculentus moench

Stomata on the cotyledons of Morphactin EMD 7301 and EMD 7311 treated seedlings may be anisocytic, anomocytic, paracytic, diacytio, transitional forms and haplocytic. In spite of this diversity, the most frequent type is anisocytic on both surfaces. The stomatal ontogeny may bo perigenous, mesogenous and mesoperigenous. Morphactin EMD 7301 and EMD 7311 increase the size of cotyledons in all concentrations accompanied by an increase in the number of stomata on the adaxial surface in 100, 150, 250 ppm of EMD 7301. The effect of morphactin is promotive to the size of stomata in all concentrations, but it retards the stomatal density and number of epidermal cells. This may be due to an increase in the size of the epidermal cells. The substrates, however, have no effect on the morphology and ontogeny of the stomata; probably they are inherent chracters.     

Epidermal studies in some Indian cultivars of Bougainvilleas

Epidermal studies in fifteen Indian cultivars of Bougainvilleas are described. The epidermal cells are polygonal isodiametric, or elongated with thick straight arched or slightly sinuous walls. Parallel culticular striations are radiating from guard cells. The mature stomata are anomocytic, paracytic and with a single subsidiary cell. The abnormal types noticed are: single guard cells with or without pores, arrested development, variously oriented contiguous stomata, cytoplasmic connections between nearby stomata and epidermal or subsidiary cells, and persistent stomatal cells. The development of anomocytic stomata is perigenous while that of the other types is mesogenous. Fifteen cultivars of Bougainvilleas are separated on the basis of bract colour, stomatal frequency and index per unit area.     

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.     

Development of Normal and Abnormal Stomata in some Araliaceae

The present investigation describes the ontogeny of normal andabnormal stomata for three species of the Araliaceae. The maturestomata are mostly anisocytic, occasionally paracytic, and rarelyanomocytic. Several aberrent patterns of stomatal developmenthave been noticed in anisocytic stomata and a few in paracyticones. The ontogeny of anisocytic and paracytic stomata conformsto the syndetocheilic or mesogenous type, while that of anomocyticis haplocheilic or perigenous.     

Epidermal Structures and Stomatal Ontogeny in Some Mimosaceae

The structure of trichomes and stomata on leaflets of 21 speciesof the Mimosaceae are described. Non-glandular trichomes inMimosa pudica are of three types: unicellular, with a roundedthick-walled base and a terminal unicellular body, and multicellular.Capitate, clavate, or cylindric, 3–6-celled glandularhairs are observed on leaflets of Mimosa pudica only. Leafletsare amphistomatic in all species except Adenanthera pavonina,Calliandra sp., Parkia biglandulosa, Pithecellobium dulce, andSamanea saman in which they are hypostomatic. Only paracyticstomata are found in Leucaena leucocephala and Mimosa pudica.In the rest stomata are of more than one type. In spite of thediversity, the most frequent type in these species is paracytic.Anisocytic stomata, in all cases, are secondarily derived fromparacytic ones by transverse or oblique wall formation in asubsidiary cell. Similarly some stomata with one subsidiarycell are also secondary derivatives of the paracytic ones becauseof one of the subsidiary cells assuming the form of an epidermalcell. The development has been traced in 14 species and thatof paracytic stomata may be mesogenous or mesoperigenous, thatof stomata with one subsidiary cell mesogenous but anomocyticstomata are ontogenetically perigenous. Occasionally a meristernoidis cut off from one of the subsidiary cells of a paracytic stoma.The organization of a stoma from such a meristemoid has beentraced.     

HELICOCYTIC AND ALLELOCYTIC STOMATA: UNRECOGNIZED PATTERNS IN THE DI COTYLEDON AE

Two new mesogenous stomatal patterns are described for the Dicotyledonae: helicocytic, with a helix of four or more subsidiary cells surrounding the guard cell pair, and allelocytic, with an alternating complex of three or more C-shaped subsidiary cells of graded sizes. The latter pattern may have guard cells developed parallel to the subsidiary cells (parallelocytic) or at right angles to them (diallelocytic). For both types, the complex development of regular sequences of mesogene subsidiary cells with particular attributes of size and arrangement provides strong probability of correlated ontogeny and adult pattern. They are related to mesogenous forms of the anisocytic, paracytic and diacytic patterns, but these latter may also be developed in different ways. The use of stomatal pattern data for taxonomic and evolutionary studies of dicote is discussed.     

Observations on developmental and structural aspects of stomata in six succulent species ofSenecio (Compositae)

The development and structure of stomata on the succulent leaves of six species ofSenecio are reported. It was found that in all the species investigated the development of stomata was of the mesogenous type and the structure of the stomata was anisocytic. The constancy in the development and structure of stomata in all the species of the genus investigated agrees with the view that a stomatal type and its developmental modes are constant at the genus level. Abnormalities such as contiguous stomata, degenerated guard cells, abortive and single guard cells are also recorded as natural phenomena.     

Stomatal Features in the Leaf of Aganosma dichotoma (Roth) K. Schum

Paracytic and anisocytic types of mature stomata are found inthe leaf of Aganosma dichotoma. Stomata with one guard cell,stomata with degenerated guard cells, and contiguous stomataare common. Stomata with arrested pore development are alsofound in certain cases. A single guard cell without any porehas not been designated as a stoma with one guard cell in thepresent investigation. Ontogeny of contiguous stomata have beentraced. Subsidiary cells are, morphologically, just like theircontiguous guard cells. Subsidiary cells may retain their shapeand contents even when their contiguous stoma becomes mature,or may change their shape and lose their contents. They mayor may not divide. Subsidiary cells form a whorl of more thantwo subsidiary cells around a stoma by their divisions. Degenerationof guard cell(s)— their contents and nuclei—havebeen traced. In certain cases guard cells divide forming morethan two guard cells associated to a single pore. Cytoplasmicconnections are found between two guard cells of nearby stomata,and between a guard cell and an epidermal cell. Near the wound,the epidermal cells over the veins become meristermatic givingrise to new epidermal cells but no meristemoid.     

Epidermal Structure and Development of Stomata in some Annonaceae

Epidermal structure and development of stomata are studied in20 species of the Annonaceae. Epidermal cells are polygonalor irregular in outline, isodiametnc or elongated with straight,arched, or sinuous anticlinal walls. Cuticular striations radiatingfrom guard cells or hair bases are noticed. Four types of trichomesand secretory cells are seen. The mature stomata are paracyticwith 2–6 subsidiary cells, non-contiguous at both poles,or contiguous at one or both the poles. The increase in numberof subsidiary cells is due to their divisions. Contiguous stomata,arrested development, single guard cell, and degeneration ofguard cells are observed. The development is either syndetocheilicor mesogenous.