Epidermal Structure and Stomatal Ontogeny in Some Eusporangiate Ferns

Cuticles, mature epidermis, and stomatal ontogeny in some speciesof the three genera of Ophioglossaceae and in Angiopteris evectawere studied. Development of stomata is perigenous in all Ophioglossaceaebut it is mesoperigenous (Piper type) in A. evecta.     

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,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.     

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.     

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.     

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.     

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.     

云南20种蕨类植物叶表皮微形态特征研究

利用光学显微镜对云南20种不同科、属的蕨类植物的叶表皮形态进行了观察和研究。结果表明：它们的叶片表面存在不同的附属物,而有的种类无附属物;20种蕨类植物共具8种气孔器类型(极细胞型、腋下细胞型、辐射状细胞型、双不等细胞型、无规则四细胞型、共环极细胞型、聚腋下细胞型和不规则型), 它们在气孔器组成上具多型现象,气孔均为下生型,分布方式及大小多样;叶表皮细胞的微形态在科属间表现出一定的差异,主要有不规则型、多边型或规则条状,表皮细胞垂周壁呈波纹状、深波状或波浪状。上述研究结果为蕨类植物的系统分类及演化提供依据。     

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     

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.     

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     

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     

Ultrastructure of stomatal development in early-divergent angiosperms reveals contrasting patterning and pre-patterning

Background and Aims

Angiosperm stomata consistently possess a pair of guard cells, but differ between taxa in the patterning and developmental origin of neighbour cells. Developmental studies of phylogenetically pivotal taxa are essential as comparative yardsticks for understanding the evolution of stomatal development.

Methods

We present a novel ultrastructural study of developing stomata in leaves of Amborella (Amborellales), Nymphaea and Cabomba (Nymphaeales), and Austrobaileya and Schisandra (Austrobaileyales), representing the three earliest-divergent lineages of extant angiosperms (the ANITA-grade).

Key Results

Alternative developmental pathways occur in early-divergent angiosperms, resulting partly from differences in pre-patterning and partly from the presence or absence of highly polarized (asymmetric) mitoses in the stomatal cell lineage. Amplifying divisions are absent from ANITA-grade taxa, indicating that ostensible similarities with the stomatal patterning of Arabidopsis are superficial. In Amborella, ‘squared’ pre-patterning occurs in intercostal regions, with groups of four protodermal cells typically arranged in a rectangle; most guard-mother cells are formed by asymmetric division of a precursor cell (the mesoperigenous condition) and are typically triangular or trapezoidal. In contrast, water-lily stomata are always perigenous (lacking asymmetric divisions). Austrobaileya has occasional ‘giant’ stomata.

Conclusions

Similar mature stomatal phenotypes can result from contrasting morphogenetic factors, although the results suggest that paracytic stomata are invariably the product of at least one asymmetric division. Loss of asymmetric divisions in stomatal development could be a significant factor in land plant evolution, with implications for the diversity of key structural and physiological pathways.     

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     

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     

Ontogeny of stomata in some Gentianaceae

The mature epidermis and development of stomata in some species of Gentianaceae, viz. Canscora deccusata, C.diffusa, Hoppea dichotoma, Enicostema littorale, Nymphoides cristatum and Gentiana pedicellata , are described. The stomatal development is shown to be trilabrate mesogenous in Canscora deccusata, C.diffusa, Enicostema littorale and Hoppea dichotoma , mesoperigenous in Gentiana and perigenous in Nymphoides. It is suggested that adoption of an aquatic habit by Nymphoides may have resulted in suppression of divisions in its perigenous stomata.     

In situ Observations of Stomatal Movements

Kappen, L., Andresen, G. and L?sch, R. 1987. In situ observationsof stomatal movements.—J. exp. Bot. 38: 126–141. A device is described by which stomatal movements in situ canbe observed and recorded continuously in light and in darkness.It is mounted in a conditioned CO₂ exchange measuring chamberso that stomatal movements can be observed whilst CO₂ exchange(photosynthesis and respiration) of the same leaf is measured.Advantages and limitations are discussed. By this method itwas shown that stomata of Vicia faba although responding inthe same direction to environmental stimuli exhibited a widerange of pore widths. Responses to changes of air humidity andof CO₂ content were clearly evident when the leaves were exposedto light. Before stomata closed due to decreasing water vapourpressure differences between leaf and air they showed a markedwidening of the pore. An inverse response occurred when watervapour pressure deficit decreased. In darkness stomata did notrespond to such changes. Key words: Stomata, leaf gas exchange, microscopic observation     

Stomatal responses of Vicia faba L. to indole acetic acid and abscisic acid

Evidence is presented that stomata in isolated epidermal peelsof Vicia faba L. open in darkness in response to the externalpresence of indole acetic acid (IAA) in the incubation medium.The effect of IAA is found to be overcome completely in thepresence of either TRIS or MES buffers. In the absence of buffer,V. faba stomata are shown to be influenced by IAA in a concentration-dependenttrend which reached a maximum at an [IAA] of 10^–3 molm^–3. Further investigations reveal that stomata in thisspecies can be shown to respond to the presence of IAA and anotherphytohormone, abscisic acid (ABA). IAA and ABA are demonstratedto be antagonistic in their effects provided the incubationconditions are suitable. The data are discussed in relationto stomatal responses of other species in different treatmentconditions. Recommendations are made with respect to standardizationof incubation media during epidermal peel experiments. Key words: Vicia faba, stomata, indole acetic acid, abscisic acid, buffers     

Epidermal Structure and Ontogeny of Stomata in Leaves of Anagallis arvensis, L.

Epidermal structure and development of stomata in leaves ofAnagallis arvensis has been described. The stomatal ontogenyis of the mesoperigenous type.     

Stomatal morphology and physiology explain varied sensitivity to abscisic acid across vascular plant lineages

Abscisic acid (ABA) can induce rapid stomatal closure in seed plants, but the action of this hormone on the stomata of fern and lycophyte species remains equivocal. Here, ABA-induced stomatal closure, signaling components, guard cell K⁺ and Ca²⁺ fluxes, vacuolar and actin cytoskeleton dynamics, and the permeability coefficient of guard cell protoplasts (P_f) were analyzed in species spanning the diversity of vascular land plants including 11 seed plants, 6 ferns, and 1 lycophyte. We found that all 11 seed plants exhibited ABA-induced stomatal closure, but the fern and lycophyte species did not. ABA-induced hydrogen peroxide elevation was observed in all species, but the signaling pathway downstream of nitric oxide production, including ion channel activation, was only observed in seed plants. In the angiosperm faba bean (Vicia faba), ABA application caused large vacuolar compartments to disaggregate, actin filaments to disintegrate into short fragments and P_f to increase. None of these changes was observed in the guard cells of the fern Matteuccia struthiopteris and lycophyte Selaginella moellendorffii treated with ABA, but a hypertonic osmotic solution did induce stomatal closure in fern and the lycophyte. Our results suggest that there is a major difference in the regulation of stomata between the fern and lycophyte plants and the seed plants. Importantly, these findings have uncovered the physiological and biophysical mechanisms that may have been responsible for the evolution of a stomatal response to ABA in the earliest seed plants.

Physiological and biophysical evidence for insensitivity of stomata to abscisic acid in ferns and lycophytes supports stomatal responsiveness to abscisic acid evolved after the divergence of ferns.