Historical development of the present classification of morphological types of stomates

There is a long-standing confusion between morphologic and ontogenetic classifications of stomates. The earliest scheme, by Vesque (1889) was proposed as basically ontogenetic, but it was soon widely applied to mature leaves. The ontogenetic distinction between haplocheilic and syndetocheilic stomates in gymnosperms, proposed by Florin (1931, 1933) soon suffered a similar fate. Continuing studies over the past half-century have shown that stomatal ontogeny is only poorly correlated with the mature morphology. Efforts to combine ontogenetic and morphologic features in a single scheme have led to classifications so complex as to be impractical. The explicitly morphological classification by Metcalfe and Chalk (1950) is the foundation for the most widely used present scheme, in which some 14 morphological types are recognized. The distinctions among these types are conceptually useful, though often arbitrary in practice.     

A new ontogenetic classification of stomatal types

A new ontogenetic classification of stomatal types is proposed which replaces the three ontogenetic types of Pant (1965) with seven new ones. The new classification clearly differentiates between the developmental involvement of the subsidiary cells and the purely structural relationship of the neighbouring cells. All known, and hypothetical, ontogenetic pathways of stomatal development can he incorporated into the new classification.
An, hitherto unknown, ontogenetic type which incorporates neighbouring, mesogene subsidiary, and perigene subsidiary cell elements into the stomatal complex is described from the fern, Polypodium vulgare L.     

Terminology and classification of stomata and stomatal development—a critical survey

The literature on terminology of stomata and stomatal development is reviewed and the terminology rationalized. The classification of developmental types and of the developing cells should not be combined with the morphological classification of mature stomatal complexes. The cells involved in the development should be distinguished on the basis of their origin and position in the developing stomatal complex, and not on the basis of their future form and appearance. It is unsound to distinguish any kind of cell only on the basis of a presumed future division by which it is replaced by its two daughter cells. Development of stomata begins with the formation of a stomatal meristemoid by an unequal division of a protodermal cell. A meristemoid may divide unequally to produce a new meristemoid and a mesogene cell. Stomatal meristemoids eventually function as guard-cell mother-cells. The adjective perigene is restricted to those cells that have arisen by divisions of protodermal cells surrounding the future stoma. The undivided cells surrounding protodermal cells should be termed agene cells, and not neighbouring cells, a term which should be restricted to morphological terminology.     

CUTICULAR ANATOMY OF ANGIOSPERM LEAVES FROM THE LOWER CRETACEOUS POTOMAC GROUP. I. ZONE I LEAVES

Angiosperm leaf cuticles from the oldest part of the Potomac Group reinforce previous paleobotanical evidence for a Cretaceous flowering plant diversification. Dated palynologically as Zone I of Brenner (Aptian?), these remains show a low structural diversity compared to later Potomac Group and modern angiosperms. All cuticle types conform to a single plan of stomatal construction that is unusual in its extraordinary plasticity: both the number of subsidiary cells and their arrangement vary greatly on a single epidermis, such that the stomata might be classified as paracytic, anomocytic, laterocytic, and intermediate. Such stomatal diversity is uncommon in extant angiosperms but is known from a few Magnoliidae. Many species possess secretory cells comparable to the oil cells of modern Magnoliidae, and a few show the bases of probable uniseriate hairs. None of the cuticle types can be assigned to a single modern family, but several show similarities with Chloranthaceae and Illiciales. These results support the concept that subclass Magnoliidae includes some of the most primitive living angiosperms.     

A new stomatal type inChenopodiaceae

A new stomatal type—paracytic mesoperigenous—which has not been separated from the paracytic mesogenous type in previous studies of theChenopodiaceae, is described. The frequent occurrence of this previously unknown paracytic mesoperigenous type in this family is demonstrated. As a result a new phylogenetic pathway between anisocytic mesoperigenous and paracytic mesogenous types may be drawn.     

The Systematic and Ecological Significance of Anatomical Characters of Leaves in the Tribe Thermopsideae (Fabaceae)

Anatomical characters of leaves of Chinese species of the tribe Thermopsideae, including 3 genera and 13 species, were compared. The results show that the differentiation of mesophyll cells, stomatal types, stomatal distribution and density, shape of epidermal cell, trichome shape and density, cuticular membrane and waxy ornamentation have important systematic and ecological significance among the genera of the tribe. With regard to leaf architecture, pollen morphology and external morphology studies, we suggest that Ammopiptanthus represents an isolated and primitive genus in the tribe and Piptanthus is closely related to Thermopsis. Although different vein-endings, stomatal types and trichomes indicate systematic differences, other characters, such as mesophyll type, stomatal distribution, trichome density etc. are the result of ecological adaptation among the genera. Results of the present study confirm our previous conclusion that two evolutionary trends exist in the tribe, the first towards xerophilization and the second towards cold-adaptation They are caused by the desertification of the central Asian area and the heave of intermountainous plateau. Ecological isolation may be a main evolutary motivation among the taxa in the tribe. We have obs'erved that stomatal density and size in the xeromorphic genus, Ammopipianthus, are not simply a result of the general trend in xerophytes to have high stomatal density correlating with the relatively small size. On the contrary, stomata in this genus are larger than those in the mesophytic genus, Piptanthus, and the density on the lower surface is much less than in the latter. However, stomatal density on the upper surface follows the trend in concordance with increasing aridity and altitute.     

八角目(Illiciales)植物花被片表皮比较形态

在光学显微镜和扫描电镜下,观察了八角科(Illiciaceae)八角属(Illicium Linn.)2组11种20个材料、五味子科(Schisandraceae)南五味子属(Kadsura Ksempf. ex Juss.)2亚属2组8种15个材料和五味子属(Schisandra Michx.)2亚属4组6种17个材料的花被片表皮形态特征。首次报道了八角目2个科（八角科和五味子科）3个属（八角属、南五味子属和五味子属）植物花被片表皮细胞的形状、分泌细胞的形状及分布、气孔器的形状及分布、花被片表面的纹饰。通过与八角目2个科3个属植物的叶表皮形态比较,发现花被片表皮气孔器外拱盖均为单层,与叶表皮气孔器外拱盖层数（常绿种类为双层和落叶种类为单层）之间没有相关性,还在五味子科中发现2个新的性状（气孔对和环列型气孔）。通过对两性花、雌花和雄花花被片表皮观察,发现花被片表皮形态与花的性别之间没有相关性。通过对八角属、南五味子属和五味子属花被片表皮比较,发现五味子属与南五味子属相比,其花被片表皮表现出更多的衍生性状;南五味子属与八角属相比,前者花被片表皮具有更多的衍生性状;而南五味子属花被片表皮形态与五味子属的相似性程度较大,支持五味子科包含南五味子属和五味子属、八角科包含八角属的观点。     

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.     

Stomatal architecture and evolution in basal angiosperms

Stomatal architecture-the number, form, and arrangement of specialized epidermal cells associated with stomatal guard cells-of 46 species of basal angiosperms representing all ANITA grade families and Chloranthaceae was investigated. Leaf clearings and cuticular preparations were examined with light microscopy, and a sample of 100 stomata from each specimen was coded for stomatal type and five other characters contributing to stomatal architecture. New stomatal types were defined, and many species were examined and illustrated for the first time. Character evolution was examined in light of the ANITA hypothesis using MacClade software. Analysis of character evolution, along with other evidence from this study and evidence from the literature on fossil angiosperms and other seed plant lineages, suggests that the ancestral condition of angiosperms can be described as anomo-stephanocytic, a system in which complexes lacking subdidiaries (anomocytic) intergrade with those having weakly differentiated subsidiaries arranged in a rosette (stephanocytic). From this ancestral condition, tangential divisions of contact cells led to the profusion of different types seen in early fossil angiosperms and Amborellaceae, Austrobaileyales, and derived Chloranthaceae, while the state in Nymphaeales is little modified. Formation of new, derived types by tangential division appears to be a recurrent theme in seed plant evolution.     

Leaf epidermal characters related with plant's passive resistance to pathogens vary among accessions of wild beans Phaseolus vulgaris var. aborigineus (Leguminosae–Phaseoleae)

We report here whether accessions of Phaseolus vulgaris var. aborigineus, collected from the southernmost part of the Andean domestication center of common bean, presented differences among leaf epidermal characters. Epidermal microcharacters such as deposition of wax crystalloids, cuticle thickness, trichome types, size and density, stomata types and size, stomatal density and index were examined on the surface of 66 leaves of 11 accessions of P. vulgaris var. aborigineus by means of light and scanning electron microscopy. Unlike the leaves of cultivated beans, those of P. vulgaris var. aborigineus were hypostomatous. Among epidermal characters only the stomatal density, stomatal index, and the number of trichomes on the abaxial leaf epidermis were different between bean accessions. The stomatal density and index of the accessions were inversely related to the altitude of the area of the wild bean collection. This is the first report describing the presence of diversity among epidermal characters (e.g. hypostomatous leaves) in P. vulgaris var. aborigineus that might contribute to plant resistance to pathogens.     

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.     

Models of steady-state and dynamic gas exchange responses to vapor pressure and light in Douglas fir (Pseudotsuga menziesii) saplings

Summary Three-dimensional empirical models were constructed, depicting the response surface of water use efficiency (WUE) of Pseudotsuga menziesii saplings in relation to different levels of both irradiance and leaf-to-air vapor pressure difference (VPD). The two models developed depict responses of (1) previous season needles during autumn, winter, and early spring and (2) current year needles during the summer. The steady-state stomatal and gas exchange responses to irradiance and VPD suggest that factors determining adaptive stomatal performance in Douglas fir are complex and may differ according to needle age, developmental stage, and season. Stomatal response to light varied seasonally, with the stomata being responsive during the summer and unresponsive during the autumn, winter, and early spring. Previous season needles exhibit higher maximum WUE, but can be less conservative in their total use of water than the more VPD-sensitive maturing needles.Observations of dynamic stomatal responses to step changes in VPD and irradiance were used to propose a simple model depicting a combined stomatal response to sudden changes in both VPD and irradiance similar to those that would occur with the passage of sunflecks in a forest canopy. Step changes in VPD caused transient stomatal movements opposite in direction to that of the final response, while step changes in irradiance resulted in movements only in the direction of the expected final response. On the basis of the model, it was hypothesized that the dynamic response to changes in VPD may serve to enhance the speed of stomatal opening and closing when changes in irradiance are rapid.     

The effect of blue light on stomatal oscillations and leaf turgor pressure in banana leaves

Stomatal oscillations are cyclic opening and closing of stomata, presumed to initiate from hydraulic mismatch between leaf water supply and transpiration rate. To test this assumption, mismatches between water supply and transpiration were induced using manipulations of vapour pressure deficit (VPD) and light spectrum in banana (Musa acuminata). Simultaneous measurements of gas exchange with changes in leaf turgor pressure were used to describe the hydraulic mismatches. An increase of VPD above a certain threshold caused stomatal oscillations with variable amplitudes. Oscillations in leaf turgor pressure were synchronized with stomatal oscillations and balanced only when transpiration equaled water supply. Surprisingly, changing the light spectrum from red and blue to red alone at constant VPD also induced stomatal oscillations – while the addition of blue (10%) to red light only ended oscillations. Blue light is known to induce stomatal opening and thus should increase the hydraulic mismatch, reduce the VPD threshold for oscillations and increase the oscillation amplitude. Unexpectedly, blue light reduced oscillation amplitude, increased VPD threshold and reduced turgor pressure loss. These results suggest that additionally, to the known effect of blue light on the hydroactive opening response of stomata, it can also effect stomatal movement by increased xylem–epidermis water supply.     

不同地区青檀叶片的解剖及表皮特征的比较研究

采用石蜡切片技术和叶表皮离析法,对我国8个省共15个居群的青檀(Pteroceltis tatarinowii Maxim.)叶片特征进行了比较和分析.研究发现:各居群的栅栏组织细胞层数多数为2层,显示出一定的稳定性,但不同居群的叶片横切面解剖特征存在显著性差异.叶片上表皮细胞有多边形和不规则形两种类型,而下表皮细胞均为不规则形.上表皮细胞垂周壁为平直形、圆形、浅波形或深波形,差异较明显,下表皮细胞垂周壁则仅有深波形和浅波形两种.叶表皮毛的分布位置在各居群间不同,而表皮毛的类型在居群间稳定,且在多数居群中表皮毛与周围细胞形成花环结构.各居群的气孔器类型均为不规则型且均仅在下表皮随机分布,但气孔器的密度、大小等特征在不同居群间存在显著性差异.生境地的气候因子对青檀叶的特征产生一定的影响.结果表明:青檀叶的解剖特征在不同居群间差异性较大,青檀叶表皮的气孔密度等特征在居群间有一定差异,而气孔器类型和表皮细胞形状等特征较稳定,有一定的分类学价值.     

Structure, Delimitation, Nomenclature and Classification of Stomata

The paper reviews stomatal types observed in 500 species of angiosperms besides those described in the literature and deals with the problems of their structure, delimitation, nomenclature and classification. In view of the varied definitions available in the literature for subsidiaries, stomatal types and, the definition and delimitations being variously interpreted by different workers, a modified definition for the subsidiaries and stomata is presented. In accordance with the international code of nomenclature for plants, the names of the stomata widely in use are retained (rule of priority). They have been presently classified as pericytic, desmocytic, paracytic, diacytic, anisocytic, anisotricytic, isotricytic, tetracytic, staurocytic, anomocytic, cyclocytic and a good number of varieties under each type are presented. These stomatal types are recognised on the basis of their structure rather than its ontogenetic pathways.     

Structure,Delimitation,Nomenclature and Classification of Stomata

The paper reviews stomatal types observed in 500 species of angiosperms besides those described in the literature and deals with the problems of their structure, delimitation, nomenclature and classification. In view of the varied definitions available in the literature for subsidiaries, stomatal types and,the definition and delimitations being variously interpreted by different workers, a modified definition for the subsidiaries and stomata is presented. In accordance with the international code of nomenclature for plants, the names of the stomata widely in use are retained (rule of priority). They have been presently classified as pericytic, desmocytic, paracytic, diacytic, anisocytic, anisotricytic, isotricytic, tetracytic,staurocytic, anomocytic, cyclocytic and a good number of varieties under each type are presented. These stomatal types are recognised on the basis of their structure rather than its ontogenetic pathways.     

Substitution of hydrazones for the 4' carbonyl on abscisic acid rendered it ineffective in a rapid stomatal–opening–inhibition bioassay

A rapid stomatal bioassay to test abscisic acid (ABA)–like inhibition of stomatal opening was elaborated. The assay was sensitive to low concentrations of ABA (>50% inhibition of opening at 0.02 μ M (+)-S-ABA) and was insensitive to 20 μ M (+)-S-ABA-1 -(methyl ester), which is consistent with previous work, With the original intent of developing anti-idiotypic antibodies, we made conjugates of ABA at the 4'carbonyl because of its accessibility and known requirements for other positions of the ABA molecule in siomatal signal transduction. We found that these conjugates lacked ABA-like activity and did not interfere with (+)-S-ABA in the stomatal bioassay, implying the importance of the 4'carbonyl. These observations cast doubt on the feasibility of using anti-idiolypic antibodies produced through the use of 4'-substituted ABA for the isolation of a guard-cell ABA receptor.