Revista Brasileira de Farmacognosia

Brazilian Journal of Pharmacognosy

Comparative anatomy of leaves of Kalanchoe
pinnata and K. crenata in sun and shade
conditions, as a support for their identification
Nattacha S. Moreira,1 Luana Beatriz S. Nascimento,1 Marcos
Vinicius Leal-Costa,3 Eliana S. Tavares*,1,2
1
Laboratório de Anatomia Vegetal, Departamento de Botânica, Instituto de Aop04512
Biologia, Universidade Federal do Rio de Janeiro, Brazil,
2
Programa de Pós-graduação em Biotecnologia Vegetal, Centro de Ciências da
Saúde, Universidade Federal do Rio de Janeiro, Brazil, Received 19 Oct 2012
3
Instituto Federal de Educação, Ciência e Tecnologia Fluminense, Campus Accepted 6 Feb 2012
Centro, Brazil.

Abstract: Kalanchoe pinnata (Lam.) Pers. and K. crenata (Andrews) Haw.,

Crassulaceae, are popularly used in the treatment of many diseases. Their biological
activities, such as anti-leishmaniasis and analgesic, can be useful in phytotherapy. Keywords:
hydathodes
Both species are often misidentified as the other, because of their similar popular leaf buds
uses and names, and the similar external morphology of the leaves. We investigated medicinal species
the existence of anatomical characters that will permit correct identification of phenolic idioblasts
the species grown in shade and in sun conditions. We also contribute with new
observations on the leaf anatomy of K. pinnata and K. crenata. Fixed (FAA70) leaves
were used, and their sections were embedded in Leica historesin. Hydathodes were
observed in both species, and for the first time were anatomically described in K.
crenata. The species showed anatomical differences in relation to the presence
of epidermal idioblasts only in K. crenata, the different pattern of distribution of
subepidermal idioblasts, and the presence of leaf buds only in K. pinnata. ISSN 0102-695X

Introduction Several biological activities have been

reported for K. pinnata and K. crenata, including anti-
The species of Crassulaceae are Crassulacean leishmaniasis (Muzitano et al., 2006), antinociceptive,
Acid Metabolism (CAM) plants. Unlike C3 and C4 plants, anti-inflammatory, and antidiabetic (Ojewole, 2005) for
CAM plants assimilate atmospheric CO2 into C4 acids at K. pinnata, and analgesic and anticonvulsant (Nguelefack
night, and subsequently fix this CO2 to the carbohydrate et al., 2006) for K. crenata. In both species, the leaf is
level during the following day (Cushman & Bohnert, the plant organ that is most used in folk medicine and in
1997). studies of biological activity.
Two species of this family, Kalanchoe pinnata Leaves are highly susceptible to environmental
(Lam.) Pers. and K. crenata (Andrews) Haw., are popularly variations, mainly light intensity. Leaves developed under
used to treat several diseases, including bronchitis and high light (sun leaves) are usually smaller and thicker,
gastritis (Moreira, et al. 2002; Medeiros et al. 2004; frequently have a higher density of stomata, a thicker
Silva et al., 2006). Besides their common uses, these epidermis and cuticle, and more developed mesophyll
species share characteristics related to leaf morphology, compared to leaves developed under low light (shade
including decussate, succulent, and glabrous leaves; ovate leaves) (Dickison, 2000; Schulze et al., 2002).
to elliptical leaf blades; and crenate margins (Hyakutake Knowledge of leaf anatomy is essential for
& Grotta, 1972; Anjoo & Kumar, 2010). Although K. the registration and quality control of herbal medicines
crenata has simple leaves and K. pinnata has simple or (Anvisa RDC No. 48/2004). Plants used in herbal
compound ones, their leaves are very similar, especially medicines may be subject to different degrees of shading
when K. pinnata has only simple leaves. Because of their during growth. Therefore, studies of medicinal plants
similarities, both species are known in Brazil as folha-da- grown under different light conditions are important
costa, saião, and coirama (Brito & Brito, 1993; Medeiros to examine photomorphogenic changes that may cause
et al. 2004; Silva et al., 2006; Joseph et al., 2011). problems with their identification (Milaneze-Gutierre et
Comparative anatomy of leaves of Kalanchoe pinnata and K. crenata in sun
and shade conditions, as a support for their identification
Nattacha S. Moreira et al.

al., 2003). Sudan III to reveal lipids (Sass, 1951), lugol for starch
Some anatomical studies have examined (Johansen, 1940), Coomassie brilliant blue for protein
K. pinnata (Jain et al. 2008; Anjoo & Kumar, 2010; (Fisher, 1968), and potassium dichromate for phenolic
Leal-Costa et al. 2010) and K. crenata (Hyakutake & compounds (Gabe, 1968). Measurements of epidermis
Grotta, 1972). However, these studies provided neither and mesophyll thickness, number of vascular bundles
information about the influence of environmental light over 1 mm, and stomatal density were made in the middle
intensity on the leaf anatomy of these species, nor a third of fourth-node leaves with an optical microscope
detailed anatomical description of them. Therefore, in (Zeiss Standard) equipped with a drawing tube. Three
view of the potential for development of herbal medicines repetitions of ten measurements were made on leaves of
from K. pinnata and K. crenata leaves, and considering different specimens. Statistical analysis was performed
the difficulties in differentiating between them based on by the GraphPad Instat 3.0 for Windows® program, using
their external morphology during the vegetative stage, this the t test (p<0.05). The photographs were taken by means
study aimed to contribute to their anatomical description of an Olympus CH30 light microscope with an attached
and to highlight distinguishing characters that are stable Olympus PM-C35B camera.
in different light conditions.
Results and Discussion
Material and Methods
Morphological differences between sun and shade plants
Plant material
Kalanchoe pinnata (Lam.) Pers. and K. crenata
Specimens of Kalanchoe pinnata (Lam.) Pers. (Andrews) Haw., Crassulaceae, plants grown in sun were
and K. crenata (Andrews) Haw., Crassulaceae, were taller, although sun plants are usually taller when grown in
obtained from the Botanical Garden of Rio de Janeiro. The shade (Taiz & Zeiger, 2009). In both species, the sun plants
voucher specimens were deposited at the Universidade had larger and thicker leaves than the shade ones (Figure
Federal do Rio de Janeiro Herbarium (RFA37525 and 1). This result differs from observations reported for sun
RFA37524, respectively). leaves regarding leaf size, which are generally smaller,
but not for leaf thickness (Esau, 1974). Kalanchoe crenata
Cultivation conditions showed rounded leaves with slightly crenate margins in
shade conditions, and elliptical leaves with pronounced
Young plants (4-5 months) were obtained from crenate margins in sun conditions. K. pinnata showed a
the Botanical Garden of Rio de Janeiro. Six specimens purple coloration on the petiole and leaf blade margins
of each species were planted in individual hard plastic only in sun conditions (Figure 1).
pots, with the same substrate and watering routine. Three
plants of each species were grown in sun and three in
shade (under a tree). Photosynthetically active radiation
(PAR) was measured monthly during the course of a year,
on sunny days, with a PAR sensor coupled to an FMS2
Hansatech fluorometer (Hansatech Instruments Ltd.,
King's Lynn, UK). The PAR intensity ranged from 413.9
to 801.3 μmol m-2 s-1 for the sun plants, and from 12.8 to
19.9 μmol m-2 s-1 for the shade plants.
Figure 1. Kalanchoe pinnata grown under sun (1a) and shade
Leaf anatomy (1b); Kalanchoe crenata grown under sun (1c) and shade (1d).

Three simple leaves from the fourth node of The plant height may have different patterns of
different specimens were fixed in FAA70 (Johansen, 1940). response according to the species’ adaptive capacity to
Leaf fragments were embedded in Leica Historesin® changes in light intensity (Muroya et al., 1997). Responses
and sectioned in a Spencer rotary microtome. Cross in leaf size may also differ between shade-tolerant and
sections were made in the proximal, middle, and distal intolerant species (Evans & Hughes, 1961; Dengler, 1980;
regions of the petiole, and in the base, middle-third and Dale, 1988).
apex of the leaf blade. The sections were stained with
toluidine blue and mounted in Entellan®. Paradermal Leaf anatomy
sections were cut mechanically and the fragments were
stained in hydroalcoholic safranin (Johansen, 1940). In K. pinnata epidermis, the anticlinal walls
Microchemical tests were performed on fresh material: were sinuous on both sides, but the sinuosity was more

Rev. Bras. Farmacogn. / Braz. J. Pharmacogn.

 Comparative anatomy of leaves of Kalanchoe pinnata and K. crenata in sun
and shade conditions, as a support for their identification
Nattacha S. Moreira et al.

pronounced on the adaxial surface (Figure 2a) than on was observed in Kalanchoe pumila by Chernetskyy &
the abaxial one, in sun leaves (Figure 2b). In K. crenata Weryszko-Chmielewska (2008), and is related to the
sun leaves, the anticlinal walls were straight to slightly ability of leaves of succulents to undergo cell division
sinuous on the adaxial surface (Figure 3a) and sinuous on long after the leaves are photosynthetically active (Ting
the abaxial surface (Figure 3b). In shade leaves of both & Gibbs, 1982).
species, the degree of sinuosity was the same on both In both species and cultivation conditions, the
surfaces (Figures 2c-d, 3c-d). abaxial surface had more stomata than the adaxial one.
Both species are amphistomatic with anisocytic The stomatal density was lower in shade plants (Table
stomata (Figures 2c, 3a). This state has also been reported 1). Jain et al. (2008) found a stomatal density of 18-20
for other species of Crassulaceae (Duarte & Zaneti, 2002; stomata/mm² in K. pinnata, but the light conditions were
Chernetskyy & Weryszko-Chmielewska, 2008). not described. According to Ting & Gibbs (1982), CAM
Kalanchoe pinnata and K. crenata have fully plants commonly have stomatal densities varying from 10
formed functional stomata, as well as others that are to 65 mm-2, as found in this study.
still differentiating (Figures 2d, 3b). The same trait In both species, the epidermis in cross-section

Figure 2. Aspects of the leaf anatomy of Kalanchoe pinnata grown in sun and shade. Paradermal sections: intercostal
region of blade (a-b) in sun leaves; (c-d) in shade leaves. Anisocytic stomata on both surfaces and in different levels
of differentiation (a,d - arrows). Cross sections of petiole (e,g) and blade (f,h-1): angular collenchyma in subepidermal
position (f-g). Homogeneous mesophyll in sun plants (h-1) and shade plants (h-2). Collateral vascular bundles (i-l)
more developed in sun plants (i,k) than in shade plants (j,l). Phenolic idioblasts (*). Legend: ac: angular collenchyma;
xy: xylem; ph: phloem.

Rev. Bras. Farmacogn. / Braz. J. Pharmacogn.

Comparative anatomy of leaves of Kalanchoe pinnata and K. crenata in sun
and shade conditions, as a support for their identification
Nattacha S. Moreira et al.

Figure 3. Aspects of the leaf anatomy of Kalanchoe crenata grown in sun and shade. Paradermal sections: intercostal
region of blade (a-b) in sun leaves; (c-d) in shade leaves. Anisocytic stomata occur on both surfaces and in different
levels of differentiation (a,b - arrows). Cross sections of petiole (e) and blade (f-l): angular collenchyma in the
subepidermal position (e-f). Homogeneous mesophyll in sun plants (g) and shade plants (h). Collateral vascular
bundles (i-l) more developed in sun plants (i,k) than in shade plants (j,l). Phenolic idioblasts*. Legend: ac: angular
collenchyma; xy: xylem; ph: phloem.

Table 1. Comparison of anatomical characters of Kalanchoe pinnata and Kalanchoe crenata sun and shade plants.
Stomatal density (stomata mm-2) Thickness of epidermis (μm) Thickness of Number of
Adaxial surface Abaxial surface Adaxial surface Abaxial surface mesophyll (μm) vascular bundles

K. pinnata Sun 23.33±2.78A,a 56.63±6.67B,a 37.23±6.57A,a 28.47±6.57B,a 1264.23±21.8a 1.77±0.63a

K. pinnata Shade 17.80±5.06 A,b
40.93±5.17 B,b
29.93±5.84 A,b
24.09±7.30 B,b
766.42±78.10 b
1.43±0.57b
K. crenata Sun 23.93±7.25 A,a
41.16±6.64 B,a
39.41±13.14 A,a
28.47±8.76 B,b
686.13±58.39 a
1.70±0.70a
K. crenata Shade 15.70±3.97A,b 30.63±8.22B,b 38.67±9.49A,a 28.47±7.30B,b 577.37±42.34b 1.30±0.50b
*Different letters indicate significantly different values (p≤0.05, n = 3). For each parameter, capital letters indicate a comparison of values between the adaxial
and abaxial surfaces; lower-case letters indicate a comparison between the values of sun and shade plants.

Rev. Bras. Farmacogn. / Braz. J. Pharmacogn.

 Comparative anatomy of leaves of Kalanchoe pinnata and K. crenata in sun
and shade conditions, as a support for their identification
Nattacha S. Moreira et al.

was uniseriate, with stomata at the same or slightly above than shade plants (Table 1), a feature generally described
the level of other epidermal cells (Figures 2e, 3e). These (Schulze et al., 2002). Kalanchoe crenata sun and shade
characteristics are shared by other crassulaceans (Duarte plants did not show significant differences in epidermal
& Zaneti, 2002; Chernetskyy & Weryszko-Chmielewska, thickness (Table 1). In both species and both cultivation
2008), except Kalanchoe daigrementiana, which has one conditions, the epidermal cells were thicker on the adaxial
to three epidermal cell layers (Balsamo & Uribe, 1988). surface.
Epidermal cells of the petiole varied little in In both species, sun plants had some angular
shape and size (Figures 2e, 3e). In the leaf blade, the collenchyma layers in the subepidermal position, in
cells were more rounded in the midrib (Figures 2f, 3f), the petiole (Figures. 2g, 3e), and in the blade midrib
and rectangular-flat in the intercostal region (Figures 2h, (Figures 2f, 3f). In shade plants, this tissue was less
3g-h). The epidermal cells of the leaf margin were larger developed, with a more restricted distribution. Below
than in the other regions (Figures 4a, c). The epidermis the collenchyma, some chlorenchyma layers followed by
was covered by a thin cuticle. ground parenchyma occurred.
K. pinnata sun plants had a thicker epidermis The mesophyll in both species was homogeneous

Figure 4. Anatomical aspects of the hydathodes and buds. Leaf cross sections of the hydathodes in K. pinnata (a,b) and
K. crenata (c,d). Tracheids (b,d - arrows) reaching the epithem (*), and a stoma located on the abaxial side (a,c - arrows).
Guttation under experimental conditions in K. pinnata (e) and K. crenata (f). Leaf cross sections of the structure of
the buds in K. pinnata (g,h). Bud cells protected by collenchyma on the abaxial surface (g - arrow). Vascular bundle
associated with the bud (h - arrow). Phenolic idioblasts (#) are related to hydathodes (d) and buds (h).

Rev. Bras. Farmacogn. / Braz. J. Pharmacogn.

Comparative anatomy of leaves of Kalanchoe pinnata and K. crenata in sun
and shade conditions, as a support for their identification
Nattacha S. Moreira et al.

and the mesophyll thickness was significantly greater collenchyma (Figures 2i-j, 3i-j). Small vascular bundles
in sun plants (Table 1), because of a larger number of frequently had associated angular collenchyma in sun
chlorenchyma layers (Figures 2h, 3g-h). The species plants (Figure 3k). In shade plants, the vascular bundles
shared some features that are commonly found in CAM were fewer (Table 1) and less developed (Figures 2l, 3l).
species: succulent leaves, thick mesophyll with large cells Hydathodes were observed in leaf blade margins
with large vacuoles, and relatively small intercellular in both species. In cross-section, they are delimited
spaces (Nelson et al., 2005). by a sheath (Figures 4a, c). The epithem is composed
K. pinnata and K. crenata had collateral of various shapes and sizes of cells in K. pinnata and
vascular bundles (Figures 2i-l, 3i-l), as observed for other generally isodiametric cells in K. crenata, without obvious
crassulaceans (Duarte & Zaneti, 2002; Chernetskyy & intercellular spaces (Figures 4b, d). Nuclei can be seen
Weryszko-Chmielewska, 2008). In the petiole and leaf by light microscopy. Tracheids are seen between epithem
midrib, the main vascular bundles had adjacent angular cells (Figure 4b, d). A stoma is located on the abaxial

Figure 5. Microchemical test. Phenolic idioblasts in K. pinnata (a-d) and K. crenata (e-h). Idioblasts in the petiole (a,e,g) and leaf
blade (b,c,d,f,h). In K. pinnata, the idioblast subepidermal layer can be observed, mainly in the petiole (a - arrow) and the midrib (b
- arrow). In the midrib and the intercostal region (c), idioblasts were more numerous near the abaxial side. Only K. crenata showed
idioblasts in the epidermis in the intercostal region (f - arrow).

Rev. Bras. Farmacogn. / Braz. J. Pharmacogn.

 Comparative anatomy of leaves of Kalanchoe pinnata and K. crenata in sun
and shade conditions, as a support for their identification
Nattacha S. Moreira et al.

surface (Figure 4a, c). only on the K. pinnata leaf margin. These differences
The hydathodes of K. pinnata and K. crenata remained constant under different lighting conditions.
differ from hydathodes of other families in having few Therefore, they can aid in anatomy-level identification
intercellular spaces between epithem cells. There are no of the species and consequently in quality control of K.
reports of hydathodes for the other species of Kalanchoe. pinnata and K. crenata herbal medicines.
The hydathodes are often related to guttation, and this
phenomemon was observed in K. pinnata and K. crenata Acknowledgements
in experimental conditions (glass cover - Figure 4e, f).
The buds of K. pinnata are located in the The authors express their gratitude to Ms.
deeper recesses of the leaf margin. They contain small Yara Lucia Oliveira de Britto, Rio de Janeiro Botanical
meristem cells with visible nuclei and more differentiated Garden, for her contribution to this study; and to Janet
isodiametric cells, some of them without an obvious W. Reid for English revision. This work was supported
nucleus. These cells are protected by collenchyma on by the Fundação de Amparo à Pesquisa do Estado do Rio
the abaxial side (Figure 4g). There is also a vascular de Janeiro and the Coordenação de Aperfeiçoamento de
bundle associated with the bud (Figure 4h). According to Pessoal de Nível Superior.
Yarbrough (1932), as the leaf expands there is a tendency
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