Stomata in some Bignoniaceae

Epidermal Structure and Ontogeny of Stomata in some Bignoniaceae

by Paliwal G. S. (1970)

In Indian Science Congress Association Proceedings 57(4): 264

(No abstract found)


The unusual leaf epidermis and stomata of Caryodaphnopsis (Lauraceae)

Origin and evolution of the unusual leaf epidermis of Caryodaphnopsis (Lauraceae)

by Zeng G., Liu B., van der Werff H., Ferguson D. K., Yang Y. (2014)

  1. Gang Zeng – Institute of Botany; University of Chinese Academy of Sciences
  2. Bing Liu – Institute of Botany; University of Chinese Academy of Sciences
  3. Henk van der Werff – Missouri Botanical Garden
  4. David K. Ferguson – University of Vienna
  5. Yong Yang – Institute of Botany; University of Chinese Academy of Sciences


In Perspectives in Plant Ecology Evolution and Systematics, 16(6), 296-309 – –


We studied leaf epidermal anatomy of Caryodaphnopsis Airy Shaw, a genus disjunct between tropical Asia and tropical America, using light microscope and scanning electron microscope. We sampled 10 species of Caryodaphnopsis and 52 species of other Lauraceous genera.

Our observations suggest that this genus possesses a unique lower leaf epidermis. Compared with other leaves of Lauraceae, this genus has an additional layer covering the lower leaf epidermis and the stomatal apparatus. The additional layer is either closed or poriferous/reticulate. The outer periclinal walls of the lower leaf epidermis protrude outside forming hollow domes or columns, and the distal endings of these domes or columns are expanded and fused, which forms the additional covering layer.

Three different types are recognized in the genus:

(1) the middle portion of the protuberances is not contracted and the distal endings are free or adnate to each other, with only limited space between the two layers;

(2) the middle portion is slightly contracted in the outer part, the distal endings are fused, with more space between the two layers;

(3) the middle portion is conspicuously contracted and elongated into columns, the distal endings are fused and it is roomy between the two layers.

The structure of the leaf lower epidermis is reconstructed and illustrated for the first time. This unusual leaf lower epidermis of Caryodaphnopsis is derived in Lauraceae and is an autapomorphic character. Outward protrusions of the outer periclinal walls forming papillate protuberances surrounding the stomatal apparatus are also found in a few other genera including Neocinnamomum, a genus closely related to Caryodaphnopsis, but the distal endings of these protrusions are not expanded and connected.

We hypothesize that the periclinal wall of the lower leaf epidermis has been gradually modified in Lauraceae, from a smooth pattern in most genera, to papillate pattern (e.g. Neocinnamomum), and to the double layered lower leaf epidermis in Caryodaphnopsis.

The origin and evolution of this unique lower epidermis might have been related to the climatic cooling and aridification since the late Eocene.

Stomata in Lauraceae

The stomatal complex of Persea borbonia

by Edwards H. H. (1990)

H. Herbert Edwards


In Botany, 68(12), 2543-2547 – –


The ultrastructure of the guard cell complex of Persea borbonia is described.

The guard cells are embedded in the midregion of the ventral wall of subsidiary cells. The outer portions of the subsidiary cell wall is extensively thickened forming a domelike structure with a large slot opening projecting above the stomatal aperture. The outer walls of epidermal cells are also extensively thickened. The fused ventral cell walls have perforations resulting in cytoplasmic continuity between guard cells.

The guard cells and subsidiary cells contain normal cytoplasmic constituents, including chloroplasts. Preliminary studies of three other Lauraceae species, Persea americana, Sassafras albidum, and Laurus nobilis, indicate a similarly structured stomatal complex.

Stomata in Quillajaceae and Surianaceae (Fabales)

FIG. 3. Quillaja saponaria, flower and inflorescence structure. – (K) A single stomata from a nectary

Floral Morphology and Development in Quillajaceae and Surianaceae (Fabales), the Species-poor Relatives of Leguminosae and Polygalaceae

by Bello Gutierrez M. A., Hawkins J. A., Rudall P. J. (2008)

Maria Angélica Bello Gutierrez, Julie A. Hawkins, Paula J. Rudall,

In Ann. Bot. 101(9): 1433, 1491-1505 – DOI: 10.1093/aob/mcn073 –


Molecular phylogenies have suggested a new circumscription for Fabales to include Leguminosae, Quillajaceae, Surianaceae and Polygalaceae. However, recent attempts to reconstruct the interfamilial relationships of the order have resulted in several alternative hypotheses, including a sister relationship between Quillajaceae and Surianaceae, the two species-poor families of Fabales. Here, floral morphology and ontogeny of these two families are investigated to explore evidence of a potential relationship between them. Floral traits are discussed with respect to early radiation in the order. Floral buds of representatives of Quillajaceae and Surianaceae were dissected and observed using light microscopy and scanning electron microscopy. Quillajaceae and Surianaceae possess some common traits, such as inflorescence morphology and perianth initiation, but development and organization of their reproductive whorls differ. In Quillaja, initiation of the diplostemonous androecium is unidirectional, overlapping with the petal primordia. In contrast, Suriana is obdiplostemonous, and floral organ initiation is simultaneous. Independent initiation of five carpels is common to both Quillaja and Suriana, but subsequent development differs; the antesepalous carpels of Quillaja become fused proximally and exhibit two rows of ovules, and in Suriana the gynoecium is apocarpous, gynobasic, with antepetalous biovulate carpels. Differences in the reproductive development and organization of Quillajaceae and Surianaceae cast doubt on their potential sister relationship. Instead, Quillaja resembles Leguminosae in some floral traits, a hypothesis not suggested by molecular-based phylogenies. Despite implicit associations of zygomorphy with species-rich clades and actinomorphy with species-poor families in Fabales, this correlation sometimes fails due to high variation in floral symmetry. Studies considering specific derived clades and reproductive biology could address more precise hypotheses of key innovation and differential diversification in the order.

Nectary stomata in Bignoniaceae

Nuptial nectary structure of Bignoniaceae from Argentina

by Rivera G. L. (2000)

In Darwiniana 38(3-4): 227-239 – License: CC BY-NC 4.0 –


Nuptial nectary characteristics were investigated in 37 taxa of Bignoniaceae.

A nuptial nectary associated to the floral axis was found in all species. Two main types can be distinguished according to their degree of development and functionality:

1) vestigial and non-secretory and

2) well-developed and secretory.

The former is characteristic of Clytostoma spp., while the latter is found in the remaining species. Two subvarieties of the secretory type of nectary can be discerned according to their position and shape:

1) annular, found in Adenocalymma, Amphilophium, Anemopaegma, Arrabidaea,Dolichandra, Eccremocarpus, Macfadyena, Melloa, Pithecoctenium, Tabebuia, and Tecoma, and

2) cylindrical, found in Argylia, Cuspidaria, Jacaranda, Mansoa, Parabignonia, Pyrostegia, and Tynnanthus.

Anatomically, two tissues are distinguished:

1) a single-layered epidermis covered by a cuticle and a variable number of stomata, and

2) a secretory tissue composed of compactly arranged parenchyma cells.

Both nectary size and nectary/ovary ratio were usually larger in lianas (Bignonieae) than in trees (Tecomeae). Nectary type proved to be consistent among species of same genus but not among genera of same tribe. Nectary features such as vascularization, presence of trichomes and nectary type were constant within the analyzed species and therefore have a reliable taxonomic value.


Stomata in Sterculiaceae and Verbenaceae

Comparative anatomical studies of the stomatal patterns of some tree species of Sterculiaceae and Verbenaceae in Nigeria

by Ajuziogu G. C., Ejeagba P. O., Nwafor F. I. 2, Ayogu V. O., Nweze A. E., Asuzu C. U., Egonu S. N. (2018)

1 Department of Plant Science and Biotechnology, University of Nigeria, Nsukka

2 Department of Pharmacognosy and Environmental Medicines, University of Nigeria, Nsukka


In Pakistan Journal of Botany 50(2) – 1518739047.pdf –


Leaf epidermal preparations of seven species within the Sterculiaceae and Verbenaceae were examined with light microscope to determine stomata patterns and other features of taxonomic importance.

Impression technique was conducted on freshly harvested leaves to reveal important foliar epidermal characters. The stomata size, index and cell wall thickness were measured following standard procedures.

Stomata were only seen in the abaxial surface of all the species, with four types observed: anomocytic, paracytic, diacytic and anisocytic. Anomocytic type was recorded for Duranta erecta, Tectonia grandis and Vitex doniana while Gmelina arborea had diacytic type.

Sterculiaceae species showed anomocytic in Cola gigantea, anisocytic in Theobroma cacao and paracytic in Cola rostrata.

Stomata sizes, numbers and epidermal cells varied across the families. Stomatal size, index and wall thickness varied greatly within the species. Variations and similarities observed in the stomatal features provide evidence of the genetic and evolutionary relationships and therefore are of taxonomic importance.

Stomata in Qualea parviflora (Vochysiaceae)

Figure 3 Qualea parviflora Mart. leaf mesophyll from Amazonian Savannah on rocky outcrops (A, D, and G), Transition Rupestrian Cerrado (B, E, and H) and from Cerradão (C, F, and I). (Cut-cuticle, Vb-vascular bundles, Pp-palisade parenchyma, Sp-spongy parenchyma,
S-stomata, and H-hypoderma). This figure is in color in the electronic version. 

Leaf anatomy of Qualea parviflora (Vochysiaceae) in three phytophysiognomies of the Mato Grosso State, Brazil

by Ariano A. P. R., da Silva I. V. (2016)

Ana Paula Ramos ARIANO,  

Ivone Vieira da SILVA,

Universidade do Estado de Mato Grosso, Faculdade de Ciências Biológicas e Agrárias, Alta Floresta-Mato Grosso, CEP: 78.580-000, Brasil.


In Acta Amaz. 46 no.2  –


Leaves have a variety of morphological and anatomical characters mainly influenced by climatic, edaphic and biotic factors. The aim of this study was to describe the anatomical leaf traits of Qualea parviflora from three phytophysiognomies. The studied phytophysiognomies were Amazon Savannah on rocky outcrops (ASR), Transition Rupestrian Cerrado (TRC), and Cerradão (CDA). Freehand sections of the leaf blade were made and stained with 0.5% astra blue and with basic fuchsin. From the adaxial and abaxial leaf surface, freehand paradermal sections were made for epidermis analysis. The Jeffrey´s method, with modifications, was used in the epidermis dissociation process. The samples from the TRC phytophysiognomy had relatively smaller ordinary epidermal cells, higher abundance of trichomes, and mesophyll with few intercellular spaces, in comparison to the other phytophysiognomies. The leaves from the ASR phytophysiognomy had higher stomatal index (SI = 21.02), and five to six layers of sclerenchyma surrounding the midrib vascular bundle. The secondary vascular bundles had thicker cell walls and the bundle sheath extended up to the epidermal tissue of both leaf sides. Leaves from the CDA phytophysiognomy had mesomorphic environmental traits, such as a thinner cuticle. It is concluded that trees from ASR and TRC phytophysiognomies have xeromorphic traits following the environmental conditions where they occur.


As folhas são órgãos vegetativos que expressam uma variedade de características morfológicas e anatômicas influenciadas, principalmente, por fatores climáticos, edáficos e bióticos. O presente estudo objetivou levantar as características anatômicas das folhas de Qualea parviflora Mart. presente em três fitofisionomias: Savana Amazônica sobre afloramentos rochosos (SAR), Cerrado Rupestre de Transição (CRT) e Cerradão (CDA). Os cortes anatômicos foram realizados a mão livre e corados com azul de astra e fucsina básica 0,5%. Para a dissociação das epidermes, foi utilizado o método de Jeffrey modificado. As amostras da fitofisionomia de CRT apresentaram células epidérmicas comuns relativamente menores, maior quantidade de tricomas e mesofilo com poucos espaços intercelulares em relação as demais fitofisionomias. As amostras da fitofisionomia de SAR apresentaram maior índice estomático (IE: 21,05), maior quantidade de esclerênquima envolvendo o feixe vascular da nervura central (5 a 6 camadas), feixes vasculares secundários com células de paredes mais espessadas, com bainha que se estende até as células epidérmicas em ambas as faces. As amostras da fitofisionomia de CDA apresentaram caracteres de ambientes mesomórficos, como cutícula mais delgada. Pôde-se concluir que os espécimes que ocorrem em SAR e CRT apresentaram características xeromórficas que estão relacionados ao ambiente de ocorrência.

Figure 1 Frontal view of the adaxial and abaxial surfaces of Qualea parviflora Mart. leaves from the Amazonian Savannah on rocky outcrops (A and D, respectively), Transition Rupestrian Cerrado (B and E, respectively), and from Cerradão (C and F, respectively). (S-stomata). This figure is in color in the electronic version. 


The leaves are hypostomatic (Figure 1D-E) and present kidney-shaped guard cells. The leaves collected in the CDA have more elongated guard cells than the leaves collected in the other phytophysiognomies. The stomata are anomocytic in all samples, but anisocytic stomata may also be found in the CDA. Both are at the same level or slightly above the other epidermal cells, and they are heterogeneously distributed in higher amounts in the ASR (stomatal index (SI) = 21.02) (Figure 1D), and in smaller amounts in TRC (SI = 17.43), as well as in the CDA specimens (SI = 19.59).


The stomata are extremely important in anatomical studies involving different environments and radiation levels, since the increased stomatal frequency in leaves exposed to high irradiance may be an important adaptive mechanisms in drier environments (Abrams and Mostoller 1995). Qualea parviflora has large stomata and they are distributed in higher amounts in the ASR samples. The leaf takes advantage of the limited time of high relative humidity in order to perform gas exchanges under xeric conditions (Medri and Lleras 1980), which might be more efficient the higher the useful stomatal area is. Stomatal movement, density, distribution, and size are species-specific features that might change in response to environmental conditions (Camargo and Marenco 2011Marencoet al. 2014). Thus, Q. parviflora presented higher amounts of stomata in the ASR samples as a strategy to increase the photosynthetic rate in higher relative humidity periods, since this phytophysiognomy has open vegetation with abundant sunlight. A distinct strategy may be observed in the TRC phytophysiognomy, where plants have less stomata, but more trichomes.