Flora 226 (2017) 47–54
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Flora
journal homepage: www.elsevier.com/locate/flora
Internal cambium and intraxylary phloem development in Ipomoea
turbinata Lag. (Convolvulaceae)
Kishore S. Rajput
∗
, Amit D. Gondaliya
Department of Botany, Faculty of Science, The Maharaja Sayajirao University of Baroda, Vadodara 390 002, India
a r t i c l e i n f o
Article history:
Received 19 July 2016
Received in revised form 5 October 2016
Accepted 3 November 2016
Edited by Alessio Papini
Available online 15 November 2016
Keywords:
Internal phloem
Morning glory
Vessel dimorphism
Variant growth
Xylem
a b s t r a c t
Strands of phloem that are present at the periphery of pith are known as intraxylary phloem. Its presence
remains restricted to a small portion of eudicots and considered as a characteristic feature for certain
families. In the present study, Ipomoea turbinata Lag. showed development of intraxylary protophloem on
adaxial tips of protoxylem from the procambial derivatives. Subsequently, additional intraxylary sieve
elements were added from the adjacent parenchyma cells that were morphologically different from
the pith cells. In thick stems, thin walled cells located between protoxylem and intraxylary protophloem
acquired meristematic characters and formed several small segments of internal cambium. Initially, these
cambial segments produced only phloem derivatives. Soon after, these segments became bidirectional
and began to produce secondary xylem centrifugally and secondary phloem centripetally. Interestingly,
in some of the samples development of secondary phloem and xylem was observed in the same direction.
The secondary xylem formed by the internal cambium was composed of wide and fibriform vessels, fibres
and axial parenchyma. Phloem possessed sieve tube elements, companion cells and parenchyma cells
while rays in both xylem and phloem were mostly uniseriate but multiseriate rays were also observed.
© 2016 Elsevier GmbH. All rights reserved.
1. Introduction
Phloem forms an important constituent of the vascular system
and plays a pivotal role in the translocation of photosynthate syn-
thesized by leaves (Pace et al., 2015). Recently transport of signaling
molecules like mRNA, macromolecules, proteins, several amino
acids and a number of unknown secondary metabolites (Golecki
et al., 1999; Zhang et al., 2010) are also ascribed to it. In most
of the eudicots and magnoliids, distribution of secondary phloem
tissue remains restricted external to the cambium. However, in
some of the plants it may be distributed/formed within secondary
xylem referred as interxylary phloem or included phloem (IAWA
Committee, 1989) while in some of the cases it may differenti-
ate at the periphery of the pith is referred as intraxylary phloem
or internal phloem (Carlquist, 2013). Beside these patterns of the
phloem distribution, family Bignoniaceae stands distinct in which
small segments of the normal cambium produce relatively less sec-
ondary xylem as compared to secondary phloem. Therefore, such
cambial variant results in the formation of four or multiple phloem
wedges (Pace et al., 2015). In the present study, Ipomoea turbinata is
∗
Corresponding author.
E-mail address: ks.rajput15@yahoo.com (K.S. Rajput).
one of the few species that show development of secondary xylem
in quantifiable amount from the internal cambium. Moreover, for-
mation of intraxylary/internal phloem is unique i.e. in some of the
samples development of intraxylary phloem and secondary xylem,
takes place on the same direction.
The occurrence of intraxylary phloem is reported in 19 fam-
ilies (Carlquist, 1988) and the family Convolvulaceae is one of
them (Carlquist and Hanson, 1991; Rajput et al., 2008, 2014).
Recently, Carlquist (2013) reviewed diversity of inter- and intraxy-
lary phloem and its functions. Further, he stated that in some
examples of intraxylary phloem, cambial activity [referred here
as internal cambium (Rajput et al., 2014)] takes place on the
inner margin of protoxylem. The similar cambium (referred as per-
imedullar cambium) has also been reported by Angyalossy et al.
(2015) in Maripa glabra from Convolvulaceae). Internal cambium
may be functionally unidirectional i.e. producing only phloem or it
may become bidirectional and produce phloem centripetally and
xylem centrifugally. Such internal cambium is reported by earlier
workers (Fukuda, 1967; Lowell and Lucansky, 1986; Carlquist and
Hanson, 1991; Terrazas et al., 2011; Carlquist, 2012; Rajput et al.,
2008, 2014). However, no exhaustive list of such species is avail-
able in which such cambial activity occurs (Carlquist, 2013). The
thick stems of Ipomoea turbinata accumulate a considerable amount
of intraxylary secondary xylem and phloem on the pith margin.
http://dx.doi.org/10.1016/j.flora.2016.11.002
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