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Comparing stomatal densities in the sun and shade

Archchana Rajmohan Department of Biological Sciences, University of Toronto

Stomata, located on the underside of leaves, allow

vascular plants to participate in gas exchange (Fig 1). It
plays a key role in photosynthesis as it allows the
exchange of carbon dioxide and oxygen to occur. But a
large amount of water is lost through the leaves in
transpiration. The amount of water loss is affected by the
size of the pore. The opening of the stomata is caused by
changes in turgor. The differences in turgor between the
guard cells and its neighbor affect the aperture of the pore
(Waggoner & Zelitch 1965).
The amount of stomata on a leaf and its width
fluctuate in accordance to light intensity, humidity and
carbon dioxide levels. For example, droughts and high
light intensities can cause stomatal densities to increase in
leaves (Shluter et al. 2002). Generally, leaves found in the
sun have more stomata, thicker mesophylls and thicker
cuticular membranes in comparison to leaves found in the
shade (Osborn & Taylor 1990). A study conducted by
Penfound (1931) found that the stomata of sunflower
plants that were growing in full sunlight had more, but
smaller, stomata per unit area that those in the shade.
In most cases, the lower epidermis has more
stomata than the upper surface (Case 2006). Theoretically,
a plant under well-watered conditions that experiences an
increase in stomatal density would have an increase
conductance for gas exchange in photosynthesis (Shluter
et al. 2002). The higher density allows for higher amounts
of water to be transpired and for more carbon dioxide to
be taken up. An increase in gas conductance can lower the
leaf temperature as the leaf can effectively engage in leaf
cooling. This is beneficial for plants in hot environments.
But if the plant is limited by water and is located in a dry
environment, this can have a negative effect on
photosynthesis and cause the plant to desiccate (Lu et al.
The purpose of this study was to observe the stomatal
density of leaves from Euonymus europaeus (Fig 2) in
two different environmental condition, sun and shade. It is
predicted that stomatal density would be higher in the
leaves found in the sun, in comparison to the leaves found
in the shade.

Materials and methods

Euonymus europaeus, a European native plant, is a
deciduous tree from the family Celastraceae (Fig 2). This
species usually grows in the full sun or in the partial
shade. This species is commonly found in old hedges and
it grows best in nutrient-abundant and salt-poor soils
(Thomas et al. 1992).

Comparing stomatal densities of

Euonymus europaeus in different
environmental conditions

Literature cited
Case, S.B. (2006) Leaf Stomata as bioindicators:
Stimulation student research. The American Biology
Teacher, 68, 88-91
Casson, S. & Gray, J.E. (2008) Influence of
environmental factors on stomatal development. New
Phytologist, 178, 9-23


Fig. 2. European
native plant,
Euonymus europaeus

The leaves from Euonymus europaeus were gathered from

the Highland Creek Ravine. E. europaeus was located
along the human-made gravel path that ran along the
ravine. A total of 30 leaves from the two environmental
condition, sun and shade, were collected. Branches that
hung over the path and were not obstructed by other
plants were deemed to be in the sun environment.
Branches that were obstructed by other plants and were
growing towards the forest were deemed to be in the
shade environment. This produced a total sample of 60
leaves. Dead leaves and leaves with hair were avoided. A
thick swath of clear nail polish was applied on the leaf
underside. Once the polish had dried, package sealing tape
was placed on the dried nail polish area. The tape was
peeled, with the impression of the leaf surface on it. The
leaf impression was observed under a dissecting
microscope under 400x power (Fig 3). The stomata was
counted in the area that appeared to have the highest
density. This was repeated for a total of three times. An
unpaired t test was run, using graphpad, to test for
significant differences between stomatal differences in the
different environments.

Fig. 3. View of
stomata under


Fig. 1. Close up of a stomata

Leaves found in the sun had higher stomatal density (Fig

4. x=30.93, SD=5.81) than leaves found in the shade
(x=18.77, SD=2.67)

Unpaired t test demonstrated that there was a

statistically significant difference between the
stomatal density between the leaves found in the
two different conditions, shade and sun (P<0.0001,
CI=9.89 to 14.50).

Mean stomatal density


Lichtenthaler, H.K., Buschmann, C., Doll, M., Fietz,

H.J., Bach, T., Kozel, U., Meier, D. & Rahmsdorf, U.
(1981) Photosynthetic activity, chloroplast
ultrastructure, and leaf characteristics of high-light and
low-light plants and of sun and shade leaves.
Photosynthesis Research, 2, 115-141



Environmental conditions
Fig. 4.This graph demonstrated that the leaves in the sun
had a higher stomatal density , on average, than leaves in
the shade.

The results demonstrated that there was a significant
difference between the stomatal density found in sun and
shade leaves. Leaves found in the sun environment had
higher stomatal density in comparison to shade leaves
(Fig 4.)
A study conducted by Lichtenthaler et al. (1981)
observed that sun leaves of Fagus sylvatica had a higher
stomata density, smaller leaf area, low water content and
was generally thicker than the leaves found in the shade.
In the study conducted by Shluter et al. (2002), the plants
grown in low light experienced a reduction in proteins
involved in photosynthesis, such as Rubisco, as the
synthesis of these proteins is stimulated by light. This
demonstrated that the low light affects photosynthesis, a
process that stomato plays an important role in. These
previously published studies and the present study help
demonstrate that plants that have high stomatal densities
have higher stomatal conductance and photosynthetic
rates. Increased density allows for larger amounts of water
to be transpired and for more carbon dioxide to be
Future studies can look into the relationship between
the physiological response stomata has to the environment
and the developmental patterns in new organs. Casson &
Gray (2008) suggest that stomatal conductance has an
influence on the responses of mature and developing

Lu, A.M., Percy, R.G., Qualset, C.O., & Zeiger, E.

(1998) Stomatal conductance predicts yields in
irrigated Pima cotton and bread wheat grown at high
temperatures. Journal of Experiemental Botany, 49,
Osborn, J.M. & Taylor, T.N. (1990) Morphological and
ultrastructural studies of plant cuticular membranes.
Botanical Gazette, 151, 465-476
Penfound, W.T. (1931) Plant anatomy as conditioned by
light intensity and soil moisture. American Journal of
Botany, 18, 558-572
Schluter, U., Muschak, M., Berger, D. & Altmann, T.
(2002) Photosynthetic performance of an Arabidopsis
mutant with elevated stomatal density (sdd1-1) under
different light regimes. Journal of Experimental
Botany, 54, 867-874
Thomas, P.A., E-Barghathi, M. & Polwart, A. (1992)
Biological Flora of the British Isles: Euonymus
europaeus L. Journal of Ecology, 99, 345-365
Waggoner, P.E. & Zelitch, I. (1965) Transpiration and
the Stomata of Leaves. American Association for the
Advancement of Science, 150, 1413-1420

I would like to thank R. Pakirathan for assisting me with the
collection of the leaves and with the painting of the leaves
with nail polish.

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