Dragon’s blood secretion and its ecological significance

Abstract

Dragon’s blood is the name given to a red exudate produced by some plant species belonging to the genera Daemonorops, Dracaena, Croton and Pterocarpus. These are endemic to various parts of the globe. It is classified as a resin or latex depending on its mode of secretion and its chemical composition, which is species specific. This red substance functions in defence and is produced (a) constitutively and stored in preformed anatomical structures, or (b) by induction in response to traumatic events, such as mechanical injury, pathogen attack or invasion by insects. Apart from its defensive role in plants, dragon’s blood is also a valuable natural resource renowned since antiquity for its diverse medicinal properties and uses in art. Despite the great importance of dragon’s blood, our knowledge of the biological basis for its secretion is still incomplete. This review summarizes recent advances in the study of the anatomical basis for its secretion, and discusses its classification and ecological function. Bringing some clarity to these issues may also help in the commercial sourcing of dragon’s blood.

Full text

REVIEW
Dragon’s blood secretion and its ecological significance
Joanna Jura-Morawiec
1
•Mirela Tulik
2
Received: 5 February 2016 / Accepted: 1 March 2016 / Published online: 19 March 2016
ÓThe Author(s) 2016. This article is published with open access at Springerlink.com
Abstract Dragon’s blood is the name given to a red
exudate produced by some plant species belonging to the
genera Daemonorops,Dracaena,Croton and Pterocarpus.
These are endemic to various parts of the globe. It is
classified as a resin or latex depending on its mode of
secretion and its chemical composition, which is species
specific. This red substance functions in defence and is
produced (a) constitutively and stored in preformed
anatomical structures, or (b) by induction in response to
traumatic events, such as mechanical injury, pathogen
attack or invasion by insects. Apart from its defensive role
in plants, dragon’s blood is also a valuable natural resource
renowned since antiquity for its diverse medicinal proper-
ties and uses in art. Despite the great importance of
dragon’s blood, our knowledge of the biological basis for
its secretion is still incomplete. This review summarizes
recent advances in the study of the anatomical basis for its
secretion, and discusses its classification and ecological
function. Bringing some clarity to these issues may also
help in the commercial sourcing of dragon’s blood.
Keywords Resin Latex Laticifers 
Constitutive defence Induced defence
Introduction
During the course of plant evolution, adaptation to biotic
and abiotic stresses is often accompanied by modification
of the organism on a range of different levels of organi-
zation that may involve changes to its morphology,
physiology and biochemistry. Secretion of resin or latex is
only one of many defence mechanisms that protect the
plant against insect invasions or pathogen attacks (Lan-
genheim 2003). Amongst the angiosperms occurs a small
group of plants called dragon’s blood trees that have the
ability to produce a red substance referred to as dragon’s
blood, which has ecological properties. This group includes
both monocotyledonous and eudicotyledonous species
belonging to the genera Daemonorops,Dracaena,Croton
and Pterocarpus (Gupta et al. 2008). Dragon’s blood trees
are endemic to various parts of the globe. The monocots
Daemonorops spp. and Dracaena spp. are native to
Southeast Asia, and to Socotra, Canary Islands, Madeira
and Morocco, China and some countries of Southeast Asia,
respectively (Roskov et al. 2015;http://e-monocot.org/),
while the eudicots Croton spp. and Pterocarpus sp. are
native to the countries of Central and South America
(Weaver 1997; Wiersema and Leon 2013; Roskov et al.
2015). A list of species of dragon trees has recently been
included in a survey by Gupta et al. (2008). Dragon’s blood
is variously classified as resin or latex, and may be pro-
duced by cells of the stem, leaves or fruit (Table 1), taking
the form of drops or chips (Balfour 1883).
Besides its great importance for the plants that produce it,
dragon’s blood is also a natural resource valuable for
Handling Editor: Michael Heethoff.
&Joanna Jura-Morawiec
j.jura@gazeta.pl
Mirela Tulik
mirela.tulik@wl.sggw.pl
1
Polish Academy of Sciences Botanical Garden - Centre for
Biological Diversity Conservation in Powsin, Prawdziwka 2,
02-973 Warsaw, Poland
2
Department of Forest Botany, Warsaw University of Life
Sciences-WULS, Nowoursynowska 159, 02-776 Warsaw,
Poland
Chemoecology (2016) 26:101–105
DOI 10.1007/s00049-016-0212-2 CHEMOECOLOGY
123
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humans. Its antiviral, antibacterial and antifungal properties
have been known since antiquity (reviewed by Gupta et al.
2008). Currently, the antioxidant properties of its extract are
used by the cosmetic industry in the production of anti-
ageing skin creams. Studies are also being conducted to
verify its anti-cancer properties (e.g. Rossi et al. 2003; Lopes
et al. 2004; Gonzalez and Valerio 2006). The increasing
demand of this natural resource has resulted in the overex-
ploitation of dragon’s blood-producing trees, and this is one
of the factors that have adversely affected the size of their
populations. Mainly, this affects Dracaena spp. By now,
both Dracaena draco and D. cinnabari, the original sources
of dragon’s blood have been included in the IUCN (Inter-
national Union for Conservation of Nature) Red List of
Threatened Species [http://www.iucnredlist.org/], and D.
cochinchinensis, which is a main source of red resin in China,
has been recognized as an endangered species in that country
(Anon 1987 after Wang et al. 2011). Although dragon’s
blood and its properties have been known since ancient
times, our knowledge of the anatomical basis for its secretion
by plants remains incomplete. One of the reasons for this
may be the restricted distribution of dragon’s blood trees.
The aim of this article is to summarize the current state of
knowledge regarding the anatomical and ecological aspects
of dragon’s blood secretion. Bringing some clarity to these
issues may also help in the commercial sourcing of dragon’s
blood.
The anatomical basis for the secretion of dragon’s
blood
Secretion and storage of resin in conifers, as well as latex
in eudicotyledonous plants, both herbaceous and tree spe-
cies, are usually associated with the formation of
specialized anatomical structures, such as resin ducts and
laticifers, respectively. In the stem of monocot arborescent
plants of the genus Dracaena, however, these special
secretory structures are absent (Fan et al. 2008). Instead,
dragon’s blood is produced by cells of the parenchymatous
ground tissue surrounding the primary and secondary
vascular bundles, as well as by cortex cells located adjacent
to the secondary protective tissue (Jura-Morawiec and
Tulik 2015). These cells have no specific morphological/
anatomical traits and can currently be identified only on the
basis of their red-coloured contents. The secretion of dra-
gon’s blood in stems of D.cochinchinensis has been
observed only in individual plants no younger than
30–50 years (Wang et al. 2010a). In D. draco, the onset of
secretion is not determined by age, and thus, dragon’s
blood is produced by young stems (Jura-Morawiec and
Tulik 2015). Red exudate has also been detected in leaf
cells of D.cochinchinensis,D.cambodiana (Wang et al.
2010b; Ou et al. 2013) and in D. draco (Fig. 1a); however,
the structural basis for its secretion has not yet been
described.
In the genus Croton, dragon’s blood is secreted by
nonarticulated laticifers and/or nonspecialized parenchyma
cells (Rudall 1987; Farias et al. 2009). The former are more
numerous in the phloem, whereas the latter are abundant in
the cortex. Laticifer abundance varies in Croton spp. and is
determined by (a) the age of the plant, (b) its position on
the tree, and (c) the environment. In general, laticifers are
less abundant in old than in young stems (Rudall 1994); the
branches have greater laticifer densities than the stem, and
trees of the tropical rain forest have more laticifers than
those of semi-deciduous tropical forests (Farias et al.
2009).
Nowadays, most dragon’s blood for commercial use is
gathered from immature fruits of rattan palms of the genus
Daemonorops (Pearson and Prendergast 2001; Baumer and
Dietemann 2010). However, neither the anatomical basis
for its secretion in this species, nor for the stem of Ptero-
carpus sp. has yet been described in the literature.
Dragon’s blood: resin, latex or sap?
According to Langenheim (2003) resin ‘‘is a lipid-soluble
mixture of volatile and non-volatile terpenoid and/or phe-
nolic secondary compounds that are usually secreted in
specialized structures located either internally or on the
Table 1 Botanical sources of dragon’s blood and its contribution to plant defence mechanism
Family Genus Species Origin Plant defence
Monocots
Arecaceae Daemonorops spp. 6 Fruit Constitutive
Asparagaceae Dracaena spp. 3 Stem, leaf Induced
Eudicots
Euphorbiaceae Croton spp. 8 Stem Constitutive
Fabaceae Pterocarpus sp. 1 Stem Constitutive
The number of species after Gupta et al. (2008)
102 J. Jura-Morawiec, M. Tulik
123
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surface of the plant and are of potential significance in
ecological interactions’’. By contrast, latex, is a mixture of
terpenoids, phenolic compounds, acids, carbohydrates, etc.
having a protective role (Lewisohn 1991) and produced in
special cells called laticifers (Fahn 1979). Chemical char-
acterization of dragon’s blood is species specific and has
been undertaken by many authors. For example, it is pos-
sible to distinguish between dragon’s blood from some
individual species used in works of art, since it has been
sold as a colourant for many centuries (Baumer and
Dietemann 2010). Dragon’s blood of Croton spp. is usually
referred to as latex due to the fact that it is secreted and
stored by laticifers, and its major constituents are poly-
meric anthocyanidins, which co-occur with many minor
constituents, including diterpenes and simple phenols
(Salatino et al. 2007). Dragon’s blood secreted by stems of
Pterocarpus officinalis is also called latex (Weaver 1997;
Guerrero and Guzman 2004); however, information about
the chemical composition of the exudate and its ecological
function is poorly known. Dragon’s blood derived from
species of Dracaena and Daemonorops is a phenolic resin
(Langenheim 2003), with well-recognized chemical con-
tent (e.g. Gonzalez et al. 2000; Shen et al. 2007; Sousa
et al. 2008). Sometimes, dragon’s blood is referred to as
sap (e.g. Philipson 2001). However, this could prove to be
a source of confusion, since plants produce other exudates
referred to by that name, such as xylem sap and phloem
sap, which are entirely different in terms of their location,
chemical composition and function. Xylem sap is trans-
ported along xylem vessels and tracheids, is mainly
composed of water, and contains several other components
such as hormones and minerals, whereas phloem sap flows
along sieve tubes (in angiosperms) and contains sugars,
amino acids, hormones and minerals dissolved in water
(Zimmermann and Brown 1971).
Red latex/resin as a plant defence strategy
Dragon’s blood may act in constitutive or induced plant
defence directed against pathogens, as well as against pests
(Table 1). As mentioned above, the chemical composition
of dragon’s blood is species specific, and is thus likely to
vary greatly with respect to the type of insects/pathogens
that may attack a given species of dragon’s blood tree. In
Croton draco, dragon’s blood is found in laticifers of the
cortex and phloem, and represents a constitutive (pre-
formed) defence. Since the bark of C. draco is very thin, it
provides little mechanical resistance, and dragon’s blood,
together with oils and tannins probably act as effective
deterrents against sucking insects and pathogens (Farias
et al. 2009). In Pterocarpus sp., dragon’s blood flows freely
when the bark is cut (Allen 1964), and therefore, one might
expect it to play a role in constitutive plant defence. Dra-
gon’s blood which coats the immature fruits of
Daemonorops spp. also seems to be a part of a constitutive
defence strategy. Lev-Yadun et al. (2009) pointed out that
the red colour of unripe fruit can serve as a warning sys-
tem, deterring herbivores from consuming chemically or
physically defended fruit, and thus, their still immature
seed. Therefore, it is likely that dragon’s blood is apose-
matic (sensu Lev-Yadun and Gould 2009; Lev-Yadun
2014). Moreover, by being sticky, it may physically and
passively entrap small organisms and, owing to its toxic
Fig. 1 Dragon’s blood of Dracaena draco.aDragon’s blood tree
growing at Jardı
´n Bota
´nico Canario ‘‘Viera y Clavijo’’ with leaves
infected by Cochinilla algodonosa, note the reddish-brown spots on
their surface (arrows). bLeaf scar (arrow) and lenticels filled with
resin on stem surface. cCross section of a lenticel. dStem wound
with margins covered with resin, the central, dead part is marked with
an asterisk.eCross-sectional view of a stem wound; the dragon’s
blood forms a barrier that isolates the infected tissue from healthy
tissues. Scale bar b,c=1mm
Dragon’s blood secretion and its ecological significance 103
123
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nature, kill them (Konno 2011). Possibly, it can also protect
against water loss and high temperatures (Langenheim
2003).
In Dracaena spp., dragon’s blood secretion can be
considered an induced natural defence mechanism, i.e.
trauma is essential for triggering its formation. This can be
due to mechanical injury, insect attack or pathogen infec-
tion (Fig. 1). Also localized tissue damage resulting from
natural developmental processes, such as leaf drop or the
formation of lenticels, may also induce dragon’s blood
secretion, as in D. draco (Fig. 1b, c). Moreover, it has been
shown that infection with the pathogenic fungi Fusarium
(Wang et al. 2010a,2011; Jiang et al. 2003), Gibberella
and Septoria (Cui et al. 2013) stimulates the production
and accumulation of resin in Dracaena spp. Following a
traumatic event, dragon’s blood is synthesized and accu-
mulates in the cells that border the wound or infected
tissues. In this way, it helps prevent the spread of the
pathogen and acts as a barrier between the wound/infected
cells and healthy tissues (Fig. 1e) (sensu Shigo 1984).
During the process of wound repair, dragon’s blood coats
the margins of the wound (Fig. 1d) providing additional
protection, possibly against desiccation. The red coloura-
tion indicating the presence of resin appears about two
weeks following the wounding of a D. draco stem (Jura-
Morawiec and Tulik 2015). This indicates that at first, the
plant relies mainly on constitutive bark defences in the
form of polyphenolic inclusions or calcium oxalate crystals
(Nagy et al. 2000; Nakata 2012), both of which have been
documented for Dracaena spp. (Prychid and Rudall 1999;
Jura-Morawiec and Tulik 2015).
Summary
Dragon’s blood secretion is a specialization occurring in a
small group of plant taxa. Its composition and mode of
secretion combine to produce effective defence mecha-
nisms that have evolved along different pathways in
species distributed across the globe. For some species, this
feature has become a double-edged sword. On the one
hand, it provides natural defence, but on the other, it makes
the plant vulnerable to exploitation. Considering that some
dragon’s blood tree species are in decline, it becomes
increasingly important that the secretion of dragon’s blood
is understood to help establish a sustainable harvest of red
resin/latex for commercial use.
Acknowledgments J. J-M thanks colleagues from Jardı
´n Bota
´nico
Canario ‘‘Viera y Clavijo’’ (Gran Canaria, Spain) for providing
information on insects that affect the leaves of Dracaena draco. This
study was funded by the Polish Academy of Sciences Botanical
Garden-Centre for Biological Diversity Conservation in Powsin under
the statutory fund.
Compliance with ethical standards
Conflict of interest The authors declare that there is no conflict of
interest.
Open Access This article is distributed under the terms of the
Creative Commons Attribution 4.0 International License (http://
creativecommons.org/licenses/by/4.0/), which permits unrestricted
use, distribution, and reproduction in any medium, provided you give
appropriate credit to the original author(s) and the source, provide a
link to the Creative Commons license, and indicate if changes were
made.
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