Stomata in selected species of Lythraceae

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Figure 1. A- Ammannia Baccifera subsp. baccifera (A1.abaxial surface with polygonal or irregular epidermal cells and wavy to sinuate anticlinal cell wall & A2. adaxial surface with polygonal epidermal cells and straight to curved anticlinal cell wall); B- Ammannia baccifera subsp. aegyptiaca (B1. abaxial surface with polygonal epidermal cells and wavy to sinuate anti clinal cell wall & B.2. adaxial surface with polygonal epidermal cells and straight to curved anticlinal cell wall); C- Ammannia multiflora (C1. abaxial & C2. adaxial surfaces- both surfaces with irregular epidermal cells and sinuate anticlinal cell wall)


Cuticular features of selected species of Ammannia, Rotala and Nesaea (Lythraceae) in South India

by Lemiya K. M., Pradeep A. K. (2017)

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in International Journal of Current Research 9(07): 54432-54440 –


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Stomata characters showed little or no variation within the clades in Lauraceae



An Evaluation of Classification By Cuticular Characters of the Lauraceae: A comparison to molecular phylogeny

by Nishida S., van der Werff H. (2011)

Sachiko Nishida, The Nagoya University Museum, Furo-cho, Chikusa-ku, Nagoya, 464-8601, Japan.


Henk van der Werff, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.



in Annals of the Missouri Botanical Garden 98(3): 348-357  –


Cuticular characters are epidermal or stomatal characters and are often used in the taxonomy and classification of fossil or extant Lauraceae. However, there is no consensus on their usefulness, especially as to which characters take priority and at which taxonomic level.

This study compared the cuticular characters of species within the Neotropical genera of the Ocotea Aubl. complex to the reported molecular phylogeny. Species of the following genera are included in this study: Aiouea Aubl., Aniba Aubl., Dicypellium Nees & Mart., Endlicheria Nees, Kubitzkia van der Werff, Licaria Aubl., Nectandra Rol. ex Rottb., OcoteaParaiaRohwer, H. G. Richt. & van der Werff, Pleurothyrium Nees, Rhodostemonodaphne Rohwer & Kubitzki, Umbellularia (Nees) Nutt., and Urbanodendron Mez.

Species groups based on cuticular characters, especially characters of the stomata, agreed well with the various clades in the molecular phylogeny, but did not agree with species grouped according to the traditional generic concepts.

Stomata characters showed little or no variation within the clades found in the molecular phylogeny. Because the number of character states is limited, cuticular features by themselves cannot be used to define genera or clades or will not allow the identification of specimens.


Stomata and a gene called MUTE

A close-up image of the surface of an Arabidopsis plant, taken under a microscope. Doughnut-shaped stomata are scattered across the surface.Soon-Ki Han/Xingyun Qi


Stomata — the plant pores that give us life — arise thanks to a gene called MUTE, scientists report

by Urton J. (2018)


in UW News May 7, 2018 –

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Close-up images of the epidermis of Arabidopsis seedlings, taken using a microscope. (A) and (C): Seedlings with typical arrangement of stomata across the surface. (B) and (D): Seedlings that artificially produce a lot of the MUTE protein, and have many stomata as a result. Scale bars are 50 micrometers.Soon-Ki Han/Xingyun Qi

Plants know how to do a neat trick.

Through photosynthesis, they use sunlight and carbon dioxide to make food, belching out the oxygen that we breathe as a byproduct. This evolutionary innovation is so central to plant identity that nearly all land plants use the same pores — called stomata — to take in carbon dioxide and release oxygen.

Without MUTE, Arabidopsis plants cannot produce stomata, and do not develop past the seedling stage.Soon-Ki Han/ Xingyun Qi

Stomata are tiny, microscopic and critical for photosynthesis. Thousands of them dot on the surface of the plants. Understanding how stomata form is critical basic information toward understanding how plants grow and produce the biomass upon which we thrive.

In a paper published May 7 in the journal Developmental Cell, a University of Washington-led team describes the delicate cellular symphony that produces tiny, functional stomata. The scientists discovered that a gene in plants known as MUTE orchestrates stomatal development. MUTE directs the activity of other genes that tell cells when to divide and not to divide — much like how a conductor tells musicians when to play and when to stay silent.

A child conducting a cellular symphony

“The MUTE gene acts as a master regulator of stomatal development,” said senior author Keiko Torii, a UW professor of biology and investigator at the Howard Hughes Medical Institute. “MUTE exerts precision control over the proper formation of stomata by initiating a single round of cell division — just one — in the precursor cell that stomata develop from.”

Stomata in Acacia (Fabaceae)

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Fig. 2. Acacia aroma. Secondary leaflets. Adx, adaxial surface; Abx, abaxial surface. ac, acroscopic semi-blade of the secondary leaflet; ba, basiscopic semi-blade of the secondary leaflet. The largest stomatal concentration is represented by the darkest shade which lightens until it is white where stomata are absent.


Stomatal distribution, stomatal density and daily leaf movement in Acacia aroma (Leguminosae)

by Hernández M. P., Arambarril A. M. (2010)

Marcelo P. Hernández1 y Ana M. ArambarriI2*

1 Cátedra de Sistemática Vegetal
2 Cátedra de Morfología Vegetal
1,2 Jardín Botánico y Arboretum ¨C. Spegazzini¨, Facultad de Ciencias Agrarias y Forestales, Universidad Nacional de La Plata, 60 y 119, C.C. 31 (1900) La Plata, Argentina 


in Boletín de la Sociedad Argentina de Botánica 45(3-4):  –


Acacia aroma Gillies ex Hook. & Arn. grows in the Chacoan and Yungas Biogeographic Provinces, Argentina. It has numerous medicinal applications, sweet and edible fruits, and it may be used as forage. The objective of the present contribution was to analyse the stomatal distribution and stomatal density on the secondary leaflet surfaces, in different parts of the leaf, and at different tree crown levels, establishing the leaf movement and environmental condition relationships. The work was performed with fresh material and herbarium specimens, using conventional anatomical techniques. Stomatal distribution on the secondary leaflet surfaces was established, and differences in stomatal density among basal, medium and apical leaflets were found. A decrease in stomatal density from the lower level to the upper level of the tree crown would be connected with that. The stomatal distribution and density appear related to the secondary leaflet shape and its position on the secondary rachis, interacting with the daily secondary leaflets and leaf movement, and the weather conditions. It is interesting that the medium value of stomata density were found in the middle part of the leaf and at the middle level of the tree crown. Original illustrations are given.


Distribución y densidad estomática y movimiento diario de la hoja en Acacia aroma (Leguminosae)Acacia aroma crece en las Provincias Biogeográficas Chaqueña y de las Yungas, Argentina. Este árbol posee numerosas aplicaciones en medicina popular, sus frutos son comestibles y puede ser usada como forraje. Los objetivos de la presente contribución fueron: establecer la distribución y densidad de los estomas en el folíolo secundario, en distintos folíolos secundarios de la misma hoja y en los folíolos secundarios de las hojas de la parte basal, media y superior de la copa del árbol, estableciendo relaciones con el movimiento diario de las hojas y condiciones ambientales. Para el estudio se utilizó material fresco y ejemplares de herbario empleando técnicas de anatomía convencionales. Se estableció la distribución de los estomas sobre las superficies adaxial y abaxial del folíolo secundario. Se encontraron diferencias en la densidad de estomas entre los folíolos secundarios de la parte basal, media y apical de la hoja que están relacionadas a la posición de éstas en la copa del árbol. Dentro de la copa del árbol se encontró que la densidad de estomas decrece desde la parte basal hasta la parte superior. La distribución y densidad estomática estarían relacionadas a la forma del folíolo secundario y posición de éstos sobre el raquis, al movimiento diario de los folíolos secundarios y de la hoja interactuando con los factores ambientales. Cabe destacar que el valor medio de densidad de estomas se halló en la parte media de la hoja y en la parte media de la copa del árbol. El trabajo se acompaña con ilustraciones originales.

Leaf anatomy and stomatal structures of six genera of Taxaceae



Leaf anatomy and its implications for phylogenetic relationships in Taxaceae s. l.

by Ghimire B., Lee C., Heo K. (2014)

Department of Applied Plant Science and Oriental Bio-herb Research Institute, Kangwon National University, Chuncheon, 200-701, Korea.


in J Plant Res. 127(3):373-388 – doi: 10.1007/s10265-014-0625-3 – Epub 2014 Feb 5 –


The comparative study on leaf anatomy and stomata structures of six genera of Taxaceae s. l. was conducted. Leaf anatomical structures were very comparable to each other in tissue shape and their arrangements. Taxus, Austrotaxus, and Pseudotaxus have no foliar resin canal, whereas Amentotaxus, Cephalotaxus, and Torreya have a single resin canal located below the vascular bundle. Among them, Torreya was unique with thick-walled, almost round sclerenchymatous epidermal cells. In addition, Amentotaxus and Torreya were comprised of some fiber cells around the vascular bundle. Also, Amentotaxus resembled Cephalotaxus harringtonia and its var. nana because they have discontinuous fibrous hypodermis. However, C. fortunei lacked the same kind of cells.

Stomata were arranged in two stomatal bands separated by a mid-vein. The most unique stomatal structure was of Taxus with papillose accessory cells forming stomatal apparatus and of Torreya with deeply seated stomata covered with a special filament structure. Some morphological and molecular studies have already been discussed for the alternative classification of taxad genera into different minor families.

The present study is also similar to these hypotheses because each genus has their own individuality in anatomical structure and stomata morphology.

In conclusion, these differences in leaf and stomata morphology neither strongly support the two tribes in Taxaceae nor fairly recognize the monogeneric family, Cephalotaxaceae. Rather, it might support an alternative classification of taxad genera in different minor families or a single family Taxaceae including Cephalotaxus.

In this study, we would prefer the latter one because there is no clear reason to separate Cephalotaxus from the rest genera of Taxaceae. Therefore, Taxaceae should be redefined with broad circumscriptions including Cephalotaxus.

PPFD and DAH are primary factors controlling stomatal function



Leaf conductance as a function of photosynthetic photon flux density and absolute humidity difference from leaf to air

Kaufmann M. R.  (1982a)

Merrill R. Kaufmann


in Plant Physiol. 69: 1018-1022 – DOI:


For an entire season of stomatal activity, leaf or needle conductance was observed on four species, each in a different genus: Engelmann spruce (Picea engelmannii Parry ex Engelm.), subalpine fir (Abies lasiocarpa [Hook.] Nutt.), lodgepole pine (Pinus contorta var. latifolia Engelm.), and aspen (Populus tremuloides Michx.).

Conductance in the natural environment was described for all species by photosynthetic photon flux density (PPFD) and absolute humidity difference from leaf to air (DAH), as follows: Conductance = b1 (√PPFD/√DAH) + b2 (√PPFD/DAH) + b3 (√PPFD/DAH2).

The only data not fitting this relationship were conifer data collected after freezing nights or aspen data collected during a short period in August when water stress occurred.

In both cases, leaf conductance was reduced. It is proposed that PPFD and DAH are primary factors controlling stomatal function for plants growing in their native range; secondary factors, such as temperature and water stress, affect conductance intermittently, except when plants are growing outside their normal environmental conditions.

Stomata in Fagus

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Figure 3. SEM micrographs of the inner surface of lower epidermis. (A) Fagus engleriana showing subsidiary cells of anomocytic type. (B) Fagus japonica showing subsidiary cells of anomocytic type. (C) Fagus longipetiolata showing subsidiary cells of anomocytic type. (D) Fagus sylvatica showing subsidiary cells of cyclocytic type. (E) Fagus hayatae showing subsidiary cells of cyclocytic type. (F) Fagus crenata showing subsidiary cells of cyclocytic type. (A–F) Scale bar = 20 μm. SEM = scanning electron microscopy.


Leaf cuticle micromorphology of Fagus L. (Fagaceae) species

by Cho S. H., Jeong K. S., Kim S.-H., Pak J.-H. (2014)

Seong Ho Cho, 12  Keum Seon Jeong, 1  Sun-Hye Kim, Jae-Hong Pak, 1

1 Department of Biology, College of Natural Sciences, Kyungpook National University, Daegu, Korea
2 Natural History Museum, Kyungpook National University, Gunwi, Korea


in Journal of Asia-Pacific Biodiversity 7(4): 378-387 –

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Figure 5. SEM micrographs of the outer surface of lower epidermis. (A) Fagus engleriana showing papillae (arrow), stomata, and well-developed waxes. (B) Fagus japonica showing trichome base (arrow) and papillae. (C) Fagus longipetiolata showing stomata and papillae (arrow). (D) Fagus crenata showing stomatal rims (arrow). (E) Fagus hayatae showing stomatal rims and weakly developed waxes. (F) Fagus sylvatica showing weakly developed waxes and stomatal rims. (A–F) Scale bar = 40 μm. SEM = scanning electron microscopy.


Cuticle micromorphology of all eight species of Fagus and an outgroup were examined in the present study. The genus Trigonobalanus was selected as the outgroup. Thirteen characteristics of the inner surface and five of the outer surface of the cuticle were described. Some characteristics, such as the subsidiary cell shape, size of stomata, arrangement of subsidiary cells, shape of anticlinal and periclinal cell walls, texture of periclinal cell wall, development epicuticular wax, and presence of papillae, were considered important for infrageneric classification.

The topology was obtained from the analysis using two major lineages: (1) Fagus engleriana + Fagus japonica + Fagus longipetiolata and (2) Fagus sylvatica + Fagus crenata + Fagus lucida + Fagus hayatae + Fagus grandifolia. The first clade supported a bootstrap value of 98% and the second clade a bootstrap value of 97%. Based on the cuticle morphology, our results support the previous study, by revealing F. englerianaF. japonica, and F. longipetiolata with long peduncles in one group and the remaining extant species of short- to medium-length peduncles in another group.

In addition, molecular phylogenetic study of Fagus based on ribosomal DNA ITS and chloroplast DNA sequences data supports this assemblage.

This study shows that cuticle micromorphological characteristics provide useful and important information for analyzing the evolutionary aspects of Fagus.


Table 2. Characters used in the cladistic analysis of Fagus and its relatives.

1. Trichome of abaxial surface: stellate & solitary (0), solitary (1), solitary and conical (2), and peltate (3)
2. Epicuticular wax of adaxial surface: weakly developed (0), developed (1), and undeveloped (2)
3. Epicuticular wax of abaxial surface: well developed (0) and weakly developed (1)
4. Papillae of abaxial surface: well developed (0) and undeveloped (1)
5. Stomatal rim: unknown (0) and well developed (1)
6. Trichome surrounding cell shape: irregular circular (0), irregular polygonal (1), and circular (2)
7. Cell number of trichome base of abaxial surface: 6–7 (0), 7–8 (1), and 10–12 (2)
8. Anticlinal wall thickness of adaxial surface: thin (0) and thick (1)
9. Anticlinal cell wall pattern of adaxial surface: straight (0), undulate (1), and sinuous and straight (2)
10. Ornament on the adaxial inner surface: absent (0) and present (1)
11. Anticlinal wall thickness of abaxial surface: thin (0) and thick (1)
12. Anticlinal cell wall pattern of abaxial surface: straight & sinuous (0), undulate (1), and sinuous (2)
13. Periclinal wall texture of abaxial surface: rough (0) and smooth (1)
14. Stomatal apparatus shape: elliptical (0), circular (1), and elliptical and circular (2)
15. Subsidiary cell number: 4–6 (0) and 6–8 (1)
16. Subsidiary cell’s anticlinal wall shape: rectangular (0), irregular circular (1), lunate (2), and circular (3)
17. Subsidiary cell arrangement: anomocytic (0), and cyclocytic and anomocytic (1)
18. Stomata length: 15–20 μm (0), 10–15 μm (1), and 20–25 μm (2)