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.     

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.     

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,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.     

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)     

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.     

Structure and Development of Stomata on the Vegetative and Floral Organs of Some Amaryllidaceae

The structure and development of stomata in 10 Amaryllidaceousmembers are described. The study is made on the leaves and floralorgans of Amaryllis belladona, Crinum asiaticum, C. bulbispermum,C. latifolium, C. pratense, Pancratium sp., Polianthes tuberosa,and Zephyranthes atamsco, and on leaves of Agave sp. and Curculigoorchioides. In most of the organs of the different plants studiedmore than one type of stoma occur, even on the same surface.However, the stomata are largely anomocytic in Amaryllis, Crinum,Pancratium, Polianthes, and Zephyranthes, but they are tetracyticin Agave and Curculigo. An increase in the number of subsidiarycells in teracytic, tricytic, paracytic, and stomata with onesubsidiary cell has been noted. It may be due to wall formationin the subsidiary cells or by the neighbouring perigenes assumingtheir form. The range of variation in different types of stomatain each organ has been studied. Abnormal features such as juxtaposedor superimposed contiguous stomata, connections between guardcells of nearby stomata, transformation of meristemoids intoepidermal cells are observed. The development of different typesof stomata in different organs of the same plant is perigenous.The systematic position of different genera is discussed inlight of the present findings. A method of preparing epidermalimprints by the use of Depex, a mounting medium, and white ofan egg is also described.     

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     

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 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     

Stomatal Structure and Stomatogenesis in Azolla pinnata R. Brown

A stoma in Azolla pinnata consists of a unicelled, binucleateguard cell with a pore, and one or more subsidiary cells towardsthe proximal side. Stomatal development is of the polocytic or mesoperigenous anomocytictype except that the guard-cell mother cell fails to form twodistinct guard cells because of restricted cytokinesis. Similaritiesand differences between the stoma of A. pinnata and that ofa fossil lycopod, Zosterophyllum myretonianum are discussed. Azolla pinnata R. Brown, Zosterophyllum myretonianum, stoma, stomatogenesis, guard cell, ferns, fossil lycopod     

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.     

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.     

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.     

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     

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     

Metabolic Inhibitors Block ABA-Induced Stomatal Closure

Closure of stomata of Commelina communis L. leaf epidermis causedby abscisic acid (ABA) was inhibited by sodium azide, potassiumcyanide and hypoxic conditions. Azide was more effective thancyanide at low concentrations, but the cyanide effect couldbe enhanced by addition of salicylhydroxamic acid, providingindirect evidence for cyanide-resistant respiration in epidermaltissue. Azide also inhibited ABA-induced closure of ‘isolated’ stomata and shrinkage of guard cell protoplasts.The results indicate that metabolic energy is required for ABAaction involving solute loss from the guard cells. Possiblemechanisms of action are discussed.     

Direct Measurements of Turgor Pressure Potentials of Guard Cells, I.

Measurements were made of pressures applied in subsidiary andguard cells which caused closure and opening of stomatal pores.These pressures were lower than would be expected from plasmolyticallydetermined osmotic potentials of guard cell saps. The pressuresneeded in guard cells to open almost closed stomata were practicallythe same as those required to close open stomata when appliedin adjacent subsidiary cells. Completely closed stomata couldnot be opened by this technique. The implications for the understandingof the mechanism of guard cell deformation, the Spannungsphase,and the pressure potentials of guard cells are discussed.     

Variable Cell Lineages form the Functional Pea Epidermis

Evidence was sought for cellular programs and cellular interactionsacting during the formation of stomatal spacing patterns. Dailyreplicas of the surfaces of Pisum sativum leaves were used toreconstruct the cellular development of specific regions ofthe epidermis. During the period studied small primordia becamemature leaves; this involved a 250-fold increase in area anda 20-fold increase in cell number. The earliest event correlatedwith the development of a stoma was an unequal division, andsuch divisions were common in neighbouring and even within thesame cells. A distinct cell lineage started with these unequaldivisions, forming both a stoma and most of the cells that separatedit from its neighbours. Both products of an unequal divisionbecame regular epidermal cells only where such development preventedthe formation of two stomata that would have been in directcontact with one another. Neighbouring stomata often developedand matured together, indicating that there was no mutual inhibitionbetween developing stomata that were more than one cell apart.It is concluded that stomata are products of an intracellularprogram which generates stomatal patterns during rather thanpreceding development. This program can be modified and evenstopped during its entire course, allowing for the correctionof local ‘mistakes’ of stomatal patterning. Cell lineages, cell determination, cellular interactions, epidermal development, garden peas, immature stomata, pattern formation, Pisum sativum, spacing patterns, stomata, unequal divisions     

Stomata on the Primary Root of Pisum sativum L.

The first record of stomata on a non-specialized root was obtainedby scanning electron microscopy of 4-d-old Pisum sativum L.In some cases subsidiary cells were trichoblasts. Stomata andthe root triarch vascular structure were simultaneously presentin transverse sections through the root. Pisum sativum, pea, root stomata, guard cells, trichoblasts