Development of stomata


Mechanisms of stomatal development: an evolutionary view

by Vatén A., Bergmann D. C. (2012)

Anne Vatén,

Dominique C. Bergmann imgres

in EvoDevo 2012, 3:11  doi:10.1186/2041-9139-3-11


Plant development has a significant postembryonic phase that is guided heavily by interactions between the plant and the outside environment. This interplay is particularly evident in the development, pattern and function of stomata, epidermal pores on the aerial surfaces of land plants. Stomata have been found in fossils dating from more than 400 million years ago. Strikingly, the morphology of the individual stomatal complex is largely unchanged, but the sizes, numbers and arrangements of stomata and their surrounding cells have diversified tremendously. In many plants, stomata arise from specialized and transient stem-cell like compartments on the leaf. Studies in the flowering plant Arabidopsis thaliana have established a basic molecular framework for the acquisition of cell fate and generation of cell polarity in these compartments, as well as describing some of the key signals and receptors required to produce stomata in organized patterns and in environmentally optimized numbers. Here we present parallel analyses of stomatal developmental pathways at morphological and molecular levels and describe the innovations made by particular clades of plants.


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Stomata of Labiatae (dicots)



by Abu-Asab M. S., Cantino P. D.

Mones S. Abu-Asab and Philip D. Cantino

in Journal of the Arnold Arboretum Vol. 68, Number 3, including pages 269-359


Because of the variety of stomatal classifications now available and the sometimes conflicting use of terms contained therein, a brief review of the situation is necessary if the reader is to understand our adopted terminology. For a more comprehensive and very enlightening review, see Rasmussen (1981). Stomata have been classified on the basis of three criteria: the configurations of neighboring and subsidiary cells in mature stomata (Vesque, 1889; Metcalfe & Chalk, 1950; Payne, 1970), stomatal ontogeny (Pant, 1965; Stevens & Martin, 1978; Payne, 1979), and a combination of the above (Fryns-Claessens & Van Cotthem, 1973; Stevens & Martin, 1978). The first criterion is relatively uncomplicated and has the advantage that it can be applied when one is working with mature leaves. Its principal disadvantage is that the same stomatal morphology may develop through different ontogenetic pathways in different plants and may therefore not be homologous (Rasmussen, 1981, and references cited therein). Classifications based partly or completely on stomatal ontogeny are more difficult to apply, and some of the terms used are defined differently by different authors. Pant (1965) classified stomata on the basis of their ontogenetic pathways: mesogenous stomata, in which the guard-cell mother cell and all subsidiaries are derived from the same meristemoid; perigenous stomata, in which all neighboring and subsidiary cells are derived from protodermal cells other than the meristemoid that produces the guard-cell mother cell; and mesoperigenous stomata, in which the surrounding cells are of dual origin, some mesogenous and others perigenous. The guard-cell mother cell is the immediate progenitor of the guard cells. Subsidiary cells surround the guard cells and clearly differ from other epidermal cells; neighboring cells immediately surround the guard cells but do not differ in shape from the remaining epidermal cells (Fryns-Claessens & Van Cotthem, 1973; Rasmussen, 1981). Unfortunately, the ambiguity of the term “meristemoid” has rendered Pant’s and other ontogenetic classifications difficult to Stomatal ontogeny starts with the unequal division of a protodermal cell. The smaller daughter cell, which contains a denser cytoplasm, divides again unequally or directly produces (by an equal division) the pair of guard cells (Fryns-Claessens & Van Cotthem, 1973; Payne, 1979; Rasmussen, 1981). The term “meristemoid” was used by Fryns-Claessens and Van Cotthem (1973) and Rasmussen (1981) to refer to the smaller daughter cell of the original protodermal cell, whereas Payne (1979) referred to the protodermal cell itself as the meristemoid. If the latter usage is adopted, there is always at least one neighboring or subsidiary cell that is derived from the meristemoid (i.e., me- sogenous), so a true perigenous type cannot exist (Fryns-Claessens & Van Cotthem, 1973; Payne, 1979). A consequent disadvantage of Payne’s terminology is that it is less precise; i.e., a wider variety of ontogenetic pathways is necessarily subsumed under the same term, mesoperigenous (see fig. 3 in Rasmussen, 1981). For this reason, and because the meristemoid sensu Payne can only be recognized after it has divided and hence no longer exists (Rasmussen, 1981), the ontogenetic terminology of Fryns-Claessens and Van Cotthem (1973) rather than that of Payne (1979) is adopted in this study. The more complex system of Stevens and Martin (1978) is even more precise but is not used here because of the difficulty in distinguishing “agene” cells (Rasmussen, 1981) from perigene cells sensu Rasmussen.

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Stomata of Euphorbia (dicots)


Anatomical studies on the genus Euphorbia L. Saudi Arabia (Subgenera: Triucalli, Ermophyton, Esula and Chamaesyce)

by Aldhebiani, A. and Jury, S.

in International Research Journal of Plant Science (ISSN: 2141-5447) Vol. 4(6) pp. 168-191, June, 2013


The genus Euphorbia is the largest in Saudi Arabia, even though no anatomical study has been done intensively.

In this study the epidermis, the stomata and the venation patterns have been investigated. The shape of the epidermal cell in Euphorbia species in Saudi Arabia varies: polygonal, rectangular, undulate or elongated. Moreover, the cell shape relies on the cell location on a leaf, i.e. the middle region, the margin, the apex or above the vein. Furthermore, in some cases both leaf surfaces have the same cell shape but more often they are unlike. Hairs are generally simple, unbranched and with a warty ornamentation on their surface. Papillae occur only in one species E. hypericifolia .

The most common stomata type is anomocytic, while the rare type is actinocytic, recorded only in E. helioscopia. Stomata of more than one type (have been encountered on the same leaf surface as in E. scordiifolia and E. hirta.

Venation patterns vary from one-veined, three-veined to those with four or more veins.


A stoma consists of the stomatal aperture and the pair of guard cells that form it. Stomata usually tend to be on the lower surface only. But in some cases this distribution varies from species to species and depends on whether the plant is a xerophyte or a mesophyte. They might be superficial or sunken. Stomata sometimes are surrounded by specialized epidermal cells which are called subsidiary cells. These subsidiaries differ from unmodified epidermal cells in shape, size and staining properties (Baranova, 1992; Metcalfe and& Chalk, 1950; Stace, 1965). On the other hand, the arrangement of subsidiary cells, where present, is of the greatest interest to the taxonomist. This variation is used to define the different types of stomata. Occasionally species have several types of stomata on one leaf, while some have only one type for the species (Stace, 1984). In addition, Van Cotthem, (1973) pointed out that those morphological stomata types can provide not only diagnostic characters but also very valuable taxonomic ones or even phylogenetic clues. Metcalfe and Chalk,(1950) had established some terms to replace the representative ‘family’ name proposed by Vesque, (1889).

Anomocytic was substituted for the ranunculaceous type; anisocytic replaced cruciferous, diacytic the caryophyllaceous and finally paracytic for the rubiaceous. The tetracytic type which can be found in most of the monocotyledons was added by Metcalfe, (1960). Later, Stace, (1965) proposed the term cyclocytic for the narrow ring of four or more subsidiary cells surrounding the stomata. Metcalfe and Chalk, (1950) have named and defined the actinocytic type as stomata surrounded by a circle of radiating cells. Three more types were introduced by Van Cotthem, (1970), hexacytic, epicytic and hemiparacytic. And some intermediate types were added by Payne, (1970) who described the helicocytic and allelocytic types in relation to mesogenous forms of anisocytic, paracytic and diacytic patterns. Stace, (1989) lists 35 types of stomata in vascular plants. Closely related families are distinguished by the presence of a specific type of stomata; such as Acanthaceae and Scrophulariaceae separated by the presence of diacytic stomata in the former as against anomocytic in the latter. Moreover, some stomatal types are distinctive of certain families: for example, Ranuculaceae has the anomocytic type, Brassicaceae the anisocytic, Caryophyllaceae diacytic, Rubiaceae paracytic and finally Poaceae has the graminaceous type (Singh, 2004). According to Metcalfe and Chalk (1950), the mature stomata of Euphorbiaceae, are anomocytic, paracytic and anisocytic. They are usually confined to the lower leaf surface, more rarely on both surfaces of the lamina. Paracytic stomata were reported by Tognini, (1897) which are mesogenous in development in E. variegata and Ricinus communis. On the other hand, according to Raju and Rao (1977), stomata in the Euphorbiaceae show considerable variation. They found that the woody taxa have predominantly the paracytic stomata type, while the anisocytic stomata are characteristic of the herbaceous Phyllanthoideae. Moreover, they indicated that Chamaesyce has a high percentage of anomocytic stomata (Raju and Rao, 1987). Finally, a considerable diversity of stomatal types were found in Euphorbia by Kakkar and Paliwal, (1974). They reported that the common type of stomata in Euphorbia species is the anomocytic, even though stomata of paracytic and anisocytic have been also observed.



Stomata of Elaphoglossum (fern)


Elaphoglossum (Dryopteridaceae-Fern) of Amazon Rainforest in Brazil: Anatomic Characterization and Adaptative Strategies

by Feio A. C., Andrade de Aguiar-Dias A. C., Conceição de Vilhena Potiguara R. (2013)

Ana Carla Feio, Ana Cristina Andrade de Aguiar-Dias, Raimunda Conceição de Vilhena Potiguara

in American Journal of Plant Sciences, 2013, 4, 1863-1871


This study describes the anatomy of sterile leaves of Elaphoglossum discolor (Kuhn) C. Chr., E. flaccidum (Fée) T. Moore and E. laminarioides (Bory ex Fée) T. Moore, the most representative species of the genus in the Ecological Park of Gunma in Pará State.

It reports the main diagnostic characters and provides new systematic data for the group. In addition, it locates the production and accumulation sites of bioactive compounds to determine possible adaptive strategies of these species in the Amazon rainforest environment.

Diagnostic structural features include stoma typology, central veins and margin forms, type of mesophyll, and the presence of sclerenchymatous sheaths in the cortex, among others. Among the bioactive compounds related to defence adaptation are phenolic compounds, which occur in all three species, and alkaloids and mucilage, which are exclusive to E. laminarioides.

Of the three species studied, E. laminarioides has features that make it the best suited to the rainforest environment.

While polocytic stomata predominated in all species, they were associated to copolocytic and anomocytic stomata in Elaphoglossum laminarioides. Although polo- cytic and copolocytic stomata are among the five types that can be found in Elaphoglossum [36], anomocytic stomata are not described for this genus. [37,38] did not correlate this stoma type with those found in E. laminarioides, but Sen and De (1992) asserted that polocytic stomata are never associated to anomocytic ones. Nevertheless, latter studies carried out by [20,21] identified ano- mocytic stomata in Argentinean species of Elaphoglossum. This variation in stomata typology is relevant since it can be used as an infrageneric diagnostic feature.

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Stomata of Polygonaceae (dicots)


Micromorphological Investigation of Foliar Anatomy of Genera Aconogonon and Bistorta of Family Polygonaceae

by Ghazalah Y., Khan M. A., Shaheen N., Hayat M. Q. (2009)


in Int. J. Agric. Biol., 11: 285–289 (2009)


Leaf epidermal studies have been carried out on six species belonging to two genera of the family Polygonaceae.

The use of light microscopy has made possible in depth to study leaf surface features such as shape of epidermal cells, stomatal pattern, their distribution on adaxial and abaxial leaf surface and trichomes types.

Epidermal cell shapes are variable but generally polygonal.

Six different stomatal patterns are reported for Aconogonon (Meisn.) Reichenb. and Bistorta Adans.

Variation among glandular and non glandular trichomes was also noted. Cyclocytic stomata are recorded first time in Aconogonon alpinum (All.) Schur.

This anatomical study has taxonomic importance, on the basis of which identification keys are prepared.

Stomata of Pteridophytes


Research Group Pteridology

Ghent University

Modern pteridophyte research at Ghent University started with the investigation of stomata types by Van Cotthem.

Subsequent micromorphological studies produced contributions on indument characters and perispore traits in various families, as well as publications on taxonomic and floristic aspects.


See: Ugent

Stomata of Eucalyptus (dicots)


Staurocytic Stomatal Complexes in Species of Monocalyptus sensu Carr and Carr (Eucalyptus, Myrtaceae)

by DJ Carr and SGM Carr

in Australian Journal of Botany 38(1) 45 – 52 (1990)

Full text doi:10.1071/BT9900045 – © CSIRO 1990


Adult leaves of the species of five groups of Eucalyptus, previously thought to be related on grounds of comparative morphology, especially of the flower, are shown to have stomatal complexes conforming to the type, rare among angiosperms, known as staurocytic.

They develop from anisocytic complexes, typical of the seedling leaves. Adults leaves of many of the species have complexes with mostly four or five subsidiary cells. The spectrum of frequencies of subsidiary cells may be characteristic of individual species.