Flora 206 (2011) 360–364
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Flora
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Dianthus caryophyllus stems and Zantedeschia aethiopica petioles/pedicels show
anatomical features indicating efficient photosynthesis
Charilaos Yiotis, George K. Psaras
∗
Section of Plant Biology, Department of Biology, University of Patras, GR 265 00 Patras, Greece
article info
Article history:
Received 16 March 2010
Accepted 23 July 2010
Keywords:
Dianthus caryophyllus
Green stems
Palisade parenchyma
Stem anatomy
Stem photosynthesis
Zantedeschia aethiopica
abstract
The fine structure of the green stem of Dianthus caryophyllus, the leaf petiole and the flower pedicel of
Zantedeschia aethiopica were studied using light and scanning electron microscopy. It was revealed that
these non-foliar plant parts of both species possess epidermis with numerous stomata. Stomatal density of
D. caryophyllus stem was found to be relatively high (79 vs 100 per mm
2
found on leaf surface). Z. aethiopica
petioles and pedicels also possess numerous stomata (17 per mm
2
), yet stomatal density was found to
be about half of that of leaves. Anatomical differences observed between petioles and pedicels were only
minor. Stems of D. caryophyllus as well as petioles and pedicels of Z. aethiopica have a chlorenchyma-
type tissue whose fine structure is quite similar to the leaf palisade chlorenchyma. Yet, the palisade of
Z. aethiopica petioles and pedicels shows a peculiar arrangement: palisade cells are arranged with their
long axis parallel to the longitudinal organ axis. Palisade tissue found in the aforementioned non-foliar
plant organs in both species shows strong red chlorophyll auto-fluorescence under epi-fluorescence
optics, consists of cells with abundant chloroplasts, possesses high percentage of intercellular spaces (13
and 20%, respectively) and its cells expose considerable part of their surface to the intercellular air. The
fine structure of this stem palisade tissue along with the abundance of functional stomata found on the
epidermis may support efficient photosynthesis.
© 2010 Elsevier GmbH. All rights reserved.
Introduction
Photosynthesis takes place in all green plant parts, i.e. plant
organs with chloroplasts. The prime sites of photosynthesis are
leaves, yet photosynthesis in non-foliar plant parts may contribute
a substantial amount to the plant’s net carbon gain under partic-
ular conditions. For example, in several desert species stems are
the primary photosynthetic organs (Gibson, 1983; Nilsen, 1995).
Moreover, green parts of flowers (Aschan and Pfanz, 2006; Aschan
et al., 2005; He and Teo, 2007; Weiss et al., 1988), fruits (Blanke and
Lenz, 1989), or even roots (Benzing et al., 1983; Hew et al., 1984;
Kitaya et al., 2002; Moreira et al., 2009) may be photosynthetically
active.
Among the alternative photosynthetic plant parts, stems
contribute significantly to the whole plant carbon budget. Photo-
synthesis in plant stems can be distinguished into four types:
(i) crassulacean-acid metabolism (CAM) photosynthesis;
(ii) corticular or bark photosynthesis, which takes place in the cor-
tex of secondary stems;
∗
Corresponding author. Tel.: +30 2610 996765; fax: +30 2610 997411.
E-mail address: g.k.psaras@upatras.gr (G.K. Psaras).
(iii) wood photosynthesis, which occurs in ray parenchyma and in
pith cells; and
(iv) photosynthesis similar to that of leaves occurring in stomata
bearing stems, simply called stem photosynthesis.
CAM photosynthesis takes place in the stems of succulent
species. Corticular photosynthesis, wood photosynthesis and pith
photosynthesis which take place in secondary thickened stems
seem to be of importance not only for the whole carbon balance
of the plant but also for alleviating potential hypoxia since perid-
erm impedes gas exchange between internal tissues and external
atmosphere (Aschan and Pfanz, 2003).
Although all four types of photosynthesis are carried out by stem
tissues, the term stem photosynthesis is usually applied to the fourth
type of photosynthesis. The case of stem photosynthesis differs
from corticular and wood photosynthesis in that the CO
2
is taken
up through abundant stomata of the stem epidermis and carbon
fixation follows the C
3
pathway (Nilsen, 1995).
Gibson (1983) analyzing the structure of mature yet lack-
ing periderm photosynthetic stems of non-succulent dicotyledons
recognized the following four basic anatomical designs on the
basis of relative position of chlorenchyma versus supporting tis-
sues: (i) cylindrical stems with continuous chlorenchyma, lacking
peripheral supporting tissues; (ii) cylindrical or angular stems hav-
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doi:10.1016/j.flora.2010.07.004