Epidermal Stomata of a Cup-Like Gall of Litsea

Photo credit : The pink cup-like galls on the lower epidermis of Litsea acuminata leaf. These galls were at the mature stage. White bar indicates 1 cm.


Observation on the Epidermal Stomata of a Cup-Like Gall of Litsea acuminata (Lauraceae)

by Yang M.-M., Huang M.-Y., Jane W.-N., Tung G. S., Chang Y.-T., Yang C.-M. (2008)

Man-Maio Yang1 , Meng-Yuan Huang2 , Wann-Neng Jane3 , G. S. Tung4 , Yung-Ta Chang2* , Chi-Ming Yang5*

1 Department of Entomology, National Chung-Hsing University Taichung, Taiwan

2 Department of Life science, National Taiwan Normal University Taipei, Taiwan

3 Institute of Plant and Microbial Bioloy, Academia Sinica Taipei, Taiwan

4 Division of Forest Protection, Taiwan Forestry Research Institute Taipei, Taiwan

5 Research Center for Biodiversity, Academia Sinica Taipei, Taiwan


in BioFormosa 43(2): 71-75 –


Screen Shot 2018-02-07 at 20.13.23
The epidermal morphology of cup-like gall at young and mature development stages on the lower epidermis of of Litsea acuminata leaf. A, lower epidermis of mature leaf; B, epidermis of young gall; C, D, E and F, epidermis of mature gall. Notes: C, crack; GC, guard cells; S, streak; T, trough. 1 and 2 indicate the first and second rip, respectively. Black bars indicate 10 μm.


Scanning electronic microscopy was used to observe the morphology and structure of the epidermis of a cup-like gall of Litsea acuminata (Bl.) Kurata leaf.

No regular stomata were found in the epidermis of galls at any growth stages and the epidermis of the gall is smoother than that of the leaf.

At the young stage, a series of little cracks, with the size of approximate 16.5×6.7 μm2 -23.3×20.0 μm2 , line up along an axis and no underneath hole could be seen in the cracks. As the gall grows older, the little cracks further rip open wider and deeper, with a size of 20.2×21.0 μm2 -24.9×17.1 μm2 , and an underneath tunnel-like network was exposed. It is unknown whether or not the tunnel-like network plays any role in the function of air exchange for photosynthesis or insect breath.


Stomata location, stomata type, cuticular ridges of stomata in Kandelia (Rhizophoraceae)

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Fig. 2. Leaf anatomical features of Kandelia. A. SEM view of the aggregated colleters of K. obovata. B. Leaf vein and the enlarged terminal tracheid of K. obovata. C. The abaxial leaf surface of K. candel, with one cork wart (the large arrow head) and stomata (the small arrow head). D. SEM view of stomata of K. obovata. E. Leaf transverse section of K. candel, showing the dorsiventral structure. F. Leaf transverse section of K. obovata, showing the semi-isobilateral structure. Co: colleters; E: epidermis; H: hypodermis; L: Leaf; P: palisade tissue; S: stipule; Sp: spongy tissue; T: terminal tracheid. A: Bar=500 µm; B: Bar=100 µm; C: Bar=250 µm; D-F: Bar=50 µm.


Morphology on Stipules and Leaves of the Mangrove Genus Kandelia (Rhizophoraceae)

by Sheue C.-R., Liu H.-Y., Yang Y.-P. (2003)

Chiou-Rong Sheue, Ho-Yih Liu, Yuen-Po Yang,

Department of Biological Sciences, National Sun Yat-sen University, Kaohsiung 804, Taiwan.


in Taiwania, 48(4): 248-258 –



The morphology of stipules and leaves of Kandelia candel (L.) Druce and K. obovata Sheue, Liu & Yong were studied and compared.

The discrepancies of anatomical features, including stomata location, stomata type, cuticular ridges of stomata, cork warts and leaf structures, among previous literatures are clarified.

Stipules have abaxial collenchyma but without sclereid ideoblast. Colleters, finger-like rod with a stalk, aggregate into a triangular shape inside the base of the stipule. Cork warts may sporadically appear on both leaf surfaces. In addition, obvolute vernation of leaves, the pattern of leaf scar and the difference of vein angles of these two species are reported.


Various leaf stomatal traits within and among black poplar

Figure 2. Stomata on adaxial (A) and abaxial (B) leaf surface, magnification 400×, ZEISS light microscope AxioVision Release 4.8.1. Estomas en la superficie adaxial (A) y abaxial (B) de la hoja, magnificación 400×, microscopio ZEISS Light AxioVision Release 4.8.1.


Leaf stomatal traits variation within and among black poplar native populations in Serbia  –

Variación de las características de los estomas de las hojas dentro y entre álamos negros de poblaciones autóctonas en Serbia

by Čortan D., Vilotić D., Šijačić-Nikolić M., Miljković D. (2017)

Dijana Čortan a *, Dragica Vilotić b , Mirjana Šijačić-Nikolić b , Danijela Miljković c

*Corresponding author: a University of Novi Sad, Faculty of Education, Department of Natural Science and management in education, Podgorička 4, Sombor, Serbia,

b University of Belgrade, Faculty of Forestry, Department of seedling, nursery and reforestation, Kneza Višeslava 1, Belgrade, Serbia.

c University of Belgrade, Institute for Biological Research “Siniša Stanković”, Department of Evolutionary Biology, Bulevar Despota Stefana 142, Belgrade, Serbia.


in BOSQUE 38(2): 337-345 – DOI: 10.4067/S0717-92002017000200011 –



Populus nigra as a keystone riparian pioneer tree species is one of the rarest and most endangered species in Europe due to the loss of its natural habitats. Genetic diversity existence is a key factor in survival of one species, and stomata as genetically controlled trait could be used for differentiation studies. With the aim of proving stomatal phenotypic variation of the four native populations of Populus nigra located on the banks of three biggest river valleys (Dunabe, Tisa and Sava) in the region of Vojvodina in northern Serbia, we examined various leaf stomatal traits (stomatal length and width, pore length and width, stomatal density, shape coefficient and stomatal and pore area).

We tested the differences of stomatal traits among populations, interindividual variability – differences among trees, the intraindividual variability, the differences between sun-exposed and shaded leaves, among leaves nested in exposition and the differences in adaxial and abaxial leaf surface. Based on mixed model ANOVA results, interpopulation variability, as statistically significant differences, observed only for stomatal pore length and shape, while all examined traits showed interindividual variability. On the intraindividual level the results showed differences for stomatal traits, except for stomatal width, stomatal shape coefficient and stomatal density regarding leaf exposure. For better understanding of how morphological and stomatal characteristics vary in black poplar populations, further studies should be necessary involving controlled environmental conditions with the aim of examining phenotypic plasticity to changing climate conditions.


Populus nigra es una especie importantes entre las pioneras de los ríos en Europa, debido a la pérdida de su hábitat natural. La existencia de diversidad genética es un factor clave en la supervivencia de una especie y los estomas, como rasgo genéticamente controlado, es usado para estudios de diferenciación. Con el objetivo de comprobar la variación fenotípica de los estomas de cuatro poblaciones autóctonas de Populus nigra en tres valles del río más grandes del norte de Serbia (Danubio, Tisa y Sava), región de Vojvodina, fueron analizados los estomas de las hojas (longitud y ancho de estomas, longitud y ancho de poros (ostíolos), densidad estomática, coeficiente de forma y área de estomas y poros). Se analizaron las diferencias de las características de los estomas entre poblaciones, la variabilidad interindividual, la variabilidad intraindividual, las diferencias entre hojas expuestas al sol y sombreadas, entre hojas anidadas en exposición y las diferencias de superficie adaxial y abaxial de las hojas. Según los resultados del modelo mixto ANDEVA, la variabilidad entre poblaciones, estadísticamente significativa, fue observada únicamente en la longitud y la forma de los poros estomáticos, mientras que todas las características analizadas mostraron variabilidad interindividual. En el nivel intraindividual, fueron observadas diferencias en las características de los estomas, excepto en su ancho, el coeficiente de forma y la densidad estomática respecto a la exposición de las hojas. Para comprender mejor cómo varían las características morfológicas y estomáticas en Populus nigra, es necesario que investigaciones futuras incluyan condiciones medioambientales controladas, para analizar la plasticidad fenotípica para el cambio de las condiciones climáticas.

Stomata in Carpinus



Leaf, stomata and trichome morphology of the species in Carpinus genus

by Chapolagh P. I., Jalali S. S., Sonboli A., Zarafshar M. (2012)




in Journ. Taxon. Biosystem. 4(10) : 11-25 –



The species of Carpinus genus are widely distributed in the Hyrcanian and Arasbaran forests. Previous researches identified the species only by leaf and seed macro-morphological traits. Leaf morphological variations in the different ecological conditions led to some problems in taxonomy of the genus.

In the current research for first time, stomata and trichome morphology were surveyed on plant collections of Noshahr Herbarium by scanning electron microscope (SEM) and light microscope (LM). Some plant samples were collected from natural sites by the authors. First, separation accuracy of Carpinus betulus, C. schuschaensis and C. oreintalis was investigated by multivariate analysis.

Extracted components of Principal Component Analysis (PCA) were highly correlated with some leaf size parameters but could not clearly separate the three groups. Discriminate analysis proved accuracy of grouping about 64.7%.Carpinus betulus had the largest dimension in stomata and trichome trait while C. orientalis had the smallest about this trait and C. schuschaensis had the medium size between of two species.

Stomata type in C. betulus was paracytic, anomocytic, and anisocytic and C. orientalis were laterocytic and C. schuschaensis was anisocytic and laterocytic.

In contrast to other species, cells of stomata located upper than epidermal cells in C. betulus. Simple unicellular trichome was determined for the genus. Although the size and dense of trichome on the leaf and petiole were different among three species, these traits were highly associated with ecological conditions.

We concluded that these traits did not have any taxonomic significant in the genus. The current research calls for seed and bract morphology as well as molecular markers to be revised.

Density of stomata is higher in bracts than in peel of dragon fruit

Photo credit : Google

Dragon Fruit Pitaya

SEM studies on the morphology of ‘white pulp’ dragon fruit

by Kammapana L., Buanong M., Techavuthiporn C., Jitareerat P., Wongs-Aree C., Yamauchi N., Srilaong V. (xxxx)

– – ISHS Acta Horticulturae 989: Southeast Asia Symposium on Quality Management in Postharvest Systems and Asia Pacific Symposium on Postharvest Quality Management of Root and Tuber Crops – 10.17660/ActaHortic.2013.989.8 –



Dragon fruit (Hylocereus undatus) is one of the new focuses for the next source of nutritional composition in Thailand. It is a good source of dietary fiber, β-carotene, magnesium, calcium, potassium mucilage, and total phenol. However, dragon fruit rapidly loses visual quality due to chlorophyll degradation of the bract or scales. Thus, the study of yellowing in the bract of dragon fruit is very important in solving the problem of loss due to yellowing. To understand the cause of bract yellowing, morphology of the fruit peel and bract were studied. At different stages of fruit development (10, 20, 30, and 40 d after anthesis), both peel and bract were sampled and observed with Scanning Electron Microscopy (SEM) at 100×. Both were evaluated for density of the stomata.

The results showed that the bract of dragon fruit at all stages of fruit development showed significantly higher density of stomata than that of the fruit peel, by about 3-4 times. As a result, water transpired via the stomata located on the bract faster than the fruit peel. Thus, bract of dragon fruit showed rapid loss of visual quality and became unacceptable. Moreover, differences in the morphology of the fruit tissue could be related to changes at the physiological level in dragon fruit at different stages of fruit development.

Stomata in Dischidia (Apocynaceae)

Photo credit : Google

Dischidia nummularia

Leaf morphology and anatomy of selected Philippine Dischidia R. Br. (Apocynaceae: Asclepiadoideae)

by Arshed M. J., Agoo E. M. (2017)

Muhammad Jefte C. Arshed1* and Dr. Esperanza Maribel Agoo2

College of Science, Graduate School 1,2, De La Salle University, 2401 Taft Avenue, Manila 0922, Philippines


Presented at the DLSU Research Congress 2017 De La Salle University, Manila, Philippines –


Screen Shot 2018-02-05 at 12.59.44
Fig. 1. Stomata of the twelve collected Dischidia species. (A) Dischidia sp. 1. (B) Dischidia sp. 2. (C) Dischidia sp. 3. (D). Dischidia sp. 4. (E). D. nummularia. (F). D. oiantha. (G.) D. Dischidia sp. 5. (H). Dischidia sp. 6. (I). D. lancifolia. (J). D. hirsuta. (K). D. platyphylla. (L). D. ruscifolia.


The epiphytic genus Dischidia R.Br. known for its close association to Hoya R.Br., remains poorly understood genus particularly in the Philippines. Though many are uniquely found in the country, many are still inadequately documented for conservation nor advance research purposes. As many discoveries are identified only by herbarium materials and out of date, particularly its morphology and anatomy aspects. So far in the Philippines, no studies had discussed specifically the leaf morphological and anatomical characteristics of these genera undermining any potential taxonomic usefulness of both attributes.

This study seeks to determine more on the morphological and anatomical features of Dischidia leaves.

For leaf stomata and epidermal structure an impression approach was done using nail polish. Leaf anatomy were studied by preparing transverse sections of the midrib by hand cut free sectioning method. The study shows that leaf morphology of the selected Philippine Dischidia agrees well to other literatures but to no avail for significant taxonomic characters. As for the anatomical area of the leaves, isopalisade and homogenous mesophyll type were determined, the first to report about the nature of Dischidia mesophyll type. For its systematic value, this mesophyll characterization is a gray feature due to its environmental susceptibility.

Stomata in Oxalis

Figure 2.12: Light microscope photographs of three examples of the smallest and largest stomata observed within southern African Oxalis species. (a) AD surface of epistomatic O. salteri MO1137, (b) AD surface of epistomatic O. magnifolia MO1524, (c) AD surface of epistomatic O. purpurea MO344, (d) AB surface of hypostomatic O. fenestrata MO1527, (e) AD surface of amphistomatic O. nortieri MO503, (f) AD surface of epistomatic O. furcillata MO56


The phylogenetic and potential functional significance of leaf anatomical and physiological traits of southern African Oxalis

Jooste M. (2015)


MSc thesis Stellenbosch University –


1.8.3 Stomata

Both the environment and evolutionary history of plants play important roles in determining and shaping plant physiology, especially in water-stressed environments (Willson et al., 2008). Stomatal traits determine water-use eciencies, gas exchange and photosynthetic rates, so it is possible to expect correlated evolution of stomatal traits and leaf anatomical traits in similar environments. However, not much research has focussed on these traits within an evolutionary context (Beerling and Kelly (1996); Brodribb et al. (2009); Dunbar-Co et al. (2009); Brodribb (2011); Haworth et al. (2011)).

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Figure 2.16: Light microscope photographs of Oxalis taxa with epistomatic leaets with additional vein-associated stomata on the AB surface, and hypostomatic leaets with additional vein-associated stomata on the AD surface. (a) AD surface of epistomatic O. cathara (MO582), (b) AD surface of epistomatic O. ava (eld), (c) AD surface of epistomatic O. psammophila (eld), (d) AB surface of O. cathara (MO582), (e) AB surface O. ava (eld), (f) AB surface O. psammophila (eld), (g) AD surface of O. compressa (MO519), (h) AD surface of O. cf. comosa (MO568), (i) AD surface of O. pes-caprae (eld), (j) AB surface of O. compressa (MO519), (k) AB surface of O. cf. comosa (MO568), (l) AB surface of O. pes-caprae (eld). Stomatal position

The position of stomata on leaets was phylogenetically signicant across southern African Oxalis clades. Taxa with hypostomatic leaets were commonly observed in the non-South African Ionoxalis clade. A recent comparative study on three South American Oxalis species also report all studied species to be hypostomatic (O. latifolia L., O. debilis, O. corniculata) (dos Reis and Alvim, 2013). O. latifolia and O. debilis are included in sect. Ionoxalis, so this data adds to our condence of stomatal placement in this section. Denton (1973) also found that North American species from the Ionoxalis clade mostly had hypostomatic leaets. AB located stomata have been described for O. carnosa (Steudle, 1983). Members of Ionoxalis, O. corniculata and O. carnosa, are phylogenetically distantly related, so this suggests that AB located stomata are widespread and ancestral in the Oxalis genus. We thus propose that AB located stomata are the ancestral state for southern African Oxalis genus. AB located stomata still characterise the species-poor Clade 2 and Clade 3, in contrast to Clade 4, which is characterised almost universally by stomata on the AD leaet surface. The major exception to AD stomata in the speciose Clade 4 is in Clade 5, the Stellata Clade of Oberlander et al. (2011), which apart from a single species (O. imbricata MO345) is hypostomatic. This implies a fairly complex history of character change for stomatal position in the lineage leading towards Clade 5. However, it must be noted that the Stellata Clade was one of the most unexpected clades retrieved by Oberlander et al. (2011), comprising a mix of morphologically dissimilar species. Recent phylogenetic results with massively increased sampling from hundreds of low-copy nuclear loci cause this clade to disintegrate, with O. imbricata being deeply embedded in Clade 4 and the remainder of the Stellata Clade resolving as sister to Clade 3 (R. Schmickl, unpublished data). Both positions are supported by morphological characters (K. Oberlander, pers comm.). These new positions imply a much simpler scenario for the evolution of stomatal position deep in the southern African Oxalis lineage, with a single, unreversed change from an ancestral hypostomatic southern African taxon to epistomaty in Clade 4. It is possible that epistomaty could serve as a key innovation for the massive Clade 4, which contains the vast majority of southern African Oxalis diversity. Taxa with amphistomatic leaets were only observed in Clade 7 and therefore this character was regarded as unique to this clade. Amphistomatic leaves frequently occur in xeric habitats (Parkhurst, 1978) and amphistomaty is regarded as an adaptation to enable the maximum conductance of a leaf. It is then possible to assume that taxa with amphistomatic leaets evolved under conditions where high conductance rates were favoured, and are still maintained, and therefore set current taxa at an advantage. An interesting association between the stomata and epidermal pavement cell types of amphistomatic 53 taxa from Clade 7 were detected, namely all taxa with amphistomatic leaets had swollen epidermal cell types (bladder cells) on both the AD and AB leaet surfaces. It is possible that the evolution of these two characters are correlated. A similar association was reported in the Aizoaceae (Bohley et al., 2015). Further research is needed to test this potential association. Stomata in crevice Stomatal position and evolution Stomatal length, density and ploidy level Stomatal complex