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Journal of Medicinal Plants Research Vol. 5(5), pp. 652-662, 4 March, 2011
Available online at http://www.academicjournals.org/JMPR
ISSN 1996-0875 ©2011 Academic Journals
Review
Why do Euphorbiaceae tick as medicinal plants? A
review of Euphorbiaceae family and its medicinal
features
Julius T. Mwine1* and Patrick Van Damme2
1Faculty of Agriculture, Uganda Martyrs University, Nkozi, P. O. Box 5498, Kampala, Uganda.
2Faculty of Bioscience Engineering, Tropical and Subtropical Laboratory of Agronomy and Ethnobotany, University of
Ghent, Coupure links 653, 9000 Ghent, Belgium.
Accepted 21 December, 2010
Euphorbiaceae is among the large flowering plant families consisting of a wide variety of vegetative
forms some of which are plants of great importance. Its classification and chemistry have of late been
subjects of interest possibly because of the wide variety of chemical composition of its members, many
of which are poisonous but useful. In this review, we have tried to demonstrate why Euphorbiaceae are
important medicinal plants. Two important issues have come up. The worldwide distribution of the
family exposes its members, to all sorts of habitats to which they must adapt, therefore inducing a large
variety of chemicals (secondary substances) that are employed for survival/defense. Succulence and
the CAM (crassulacean acid metabolism) pathway that characterize a good number of its members were
quoted as some of the adaptations that aid colonization and survival to achieve this induction. We have
also found out that medicinal properties of some of its species may be due to stress factors that
characterize most habitats of the family. Varying stress factors like temperature, salinity, drought and
others were seen to operate in tandem with genetic factors such as gene expression and mutation
loads to bring about synthesis of a wide assemblage of secondary substances that may probably be
responsible for the family’s medicinal nature. It was concluded that the family is a good starting point
for the search for plant-based medicines.
Key words: Bio-synthesis, ethnomedicine, secondary metabolites, stress physiology.
INTRODUCTION
Family Euphorbiaceae as traditionally delimited
(Euphorbiaceae s.l., Webster 1994) is one of the largest
families of flowering plants, composed of over 300
genera and 8,000 species. According to the latter author,
the family is very diverse in range, composed of all sorts
of plants ranging from large woody trees through climbing
lianas to simple weeds that grow prostrate to the ground.
Members are widely distributed all around the world
constituting both old world and new world plants some of
which are yet to be identified. Many family members are
inhabitants of tropical climates surviving hot dry desert
conditions while others are rainforest trees and herbs.
*Corresponding author. mwinej@yahoo.com
The family consists of species of great economic
importance like Ricinus communis L. (castor oil plant),
Manihot esculenta Crantz (cassava) and Hevea
brasiliensis Willd. Ex. A. Juss (rubber tree) among others
but also noxious weeds like Euphorbia esula L. and
Euphorbia maculata L. (Schultes, 1987). The implication
of this is that Euphorbiaceae is a complex family with a
lot of research potential.
Complexity in habitat range and variability in
morphology and genetics has made Euphorbiaceae
classification difficult. Homogenous families have their
classification based on simple, unique characteristics that
cut across the family, e.g. monocotyledony and parallel
venation for family Poaceae. In the case of
Euphorbiaceae, there appears to be no particular and
easily observable feature that can be used for its
classification. In agreement with this, Webster (1994)
states that no single feature characterizes the
Euphorbiaceae. Instead, he enumerates several
anatomic features like wood structure, laticifer type,
trichomes and nature of stomata as being important for
family classification, while others like pollen nuclear
numbers, exine structures, type of pollination and
inflorescence types are important for classifying genera,
tribes and subfamilies.
According to Webster (1975), there have been several
Euphorbiaceae classifications dating as far back as 1824
by taxonomists like Adrien Jussieu who identified the
family’s genera and Jean Mueller who provided a first
detailed classification of the family into subfamilies, tribes
and sub-tribes. Webster (1975) believes that Mueller’s
classification of 1866 was a milestone in Euphorbiaceae
classification. He argues that Mueller was the first to use
coherent phylogenetic characteristics that for a long
period withstood the test of time.
Using Mueller’s classification as a skeleton, Webster
employed phylogenetic structures such as pollen
morphology and anatomy to come up with his own
classification. He divided the family into five subfamilies
that is, Acalyphoideae, Crotonoideae, Euphorbioideae,
Phyllanthoideae and Oldfieldoiideae (Webster, 1975).
According to this classification, the first three subfamilies
are characterized by one ovule per locule (uni-ovulate)
while the last two have two ovules (bi-ovulate). For
several decades, this was the (traditional and) generally
accepted form of classification.
As is common with such large and diverse families like
Euphorbiaceae, there was constant pressure and
proposals to re-define the family boundaries, to exclude
genera that appear il-fitting and include those that appear
left out, but also to carry out internal re-organization of
subfamilies, tribes and sub-tribes. Phytochemical and
molecular phylogenetic studies eventually accumulated
evidence pointing to non-monophyly of Euphorbiaceae
(Seigler, 1994b; Tokuoka and Tobe, 1995). This
culminated into partitioning of the traditional
Euphorbiaceae into five families, where only uni-ovulate
subfamilies constituted family Euphorbiaceae sensu lato,
others being upgraded with additions or subtractions into
their own families (Webster, 1994) and was validated by
the APG II group (Wurdack et al., 2005).
The new classification left family Euphorbiaceae s.l.
with five subfamilies, 49 tribes, 317 genera and about
8,000 species (Webster, 1994). However, little
experimental support met this classification and more
studies both in support and against or to further re-
organize the family have since been in progress (Bruyns
et al., 2006; Davis et al., 2007; Henderson, 1992;
Hoffmann and McPherson, 2007; Seigler, 1994; Tokuoka,
2007; Wurdack et al., 2005).
Using recent molecular study results based on DNA
sequencing with molecular markers rbcL, atpB, matK and
18S rDNA; plastids RBCL and TRNL-F and a nuclear
Mwine and Van Damme 653
gene PHYC (Tokuoka, 2007; Wurdack et al., 2005),
Euphorbiaceae s.l. family has recently been split again
into five families namely Euphorbiaceae sensu stricto,
Pandaceae, Phyllanthaceae, Picrodendraceae, and
Putranjivaceae (Tokuoka, 2007). Work on re-organization
and proof of monophyly within these groups has been
going on (Riina et al., 2010; Sierra et al., 2010;
Vorontsova and Hoffmann, 2008; Vorontsova et al., 2007;
Wurdack and Davis, 2009). According to Stevens (2010),
Euphorbiaceae Jussieu s.s is now made up of 218
genera and 5,735 species and has been subdivided into
four (supported) clades namely Chelosiodeae K.
Wurdack and Petra Hoffmann, Acalyphoideae Beilscmeid
s.s, Crotonoideae Beilschmeid s.s, and Euphorbiodeae
Beilscmeid s.s.
A note on Euphorbiaceae ethnomedicine
Just like the complexity in classification, ethnomedicine of
Euphorbiaceae is very diverse. According to Seigler
(1994), this diversity is due to the presence of a wide
range of unusual secondary metabolites that makes most
of the members poisonous. The family hosts one of the
most poisonous substances of plant origin that is, ricin,
which is a protein found in Ricinus communis (Palatnick
and Tenenbein, 2000), whereas other species like
Jatropha curcas L. are reported to be comparatively
poisonous (Mampane et al., 1987).
In an attempt to reveal the wide diversity of poisons of
the family, Abdel-Fattah (1987) lists examples of species
with following features: fish poisons e.g. Euphorbia
scheffleri Pax, Euphorbia tirucalli L., and Euphorbia
inaequilatera Sond; human poisons: Euphorbia ledienii A.
Berger, Euphorbia heterophylla L., Euphorbia cooperi
N.E.Br. ex A. Berger, Euphorbia candelabrum Kotschy,
Euphorbia virosa Willd., Euphorbia poissonii Pax,
Euphorbia unispina N.E.Br. and Euphorbia venenifica
Tremaux ex Kotschy; poisons of domestic animals:
Euphorbia caput-medusae L., Euphorbia silenifolia
(Haworth) Sweet, Euphorbia ingens E. Mey. Ex Boiss; as
well as irritating ones: E. tirucalli, Euphorbia poissonii,
Euphorbia unispina and E. venenifica. In addition, some
members are said to cause or influence susceptibility to
certain body ailments. For example E. tirucalli, Euphorbia
leuconeura, J. curcas and others are known to be co-
carcinogenic and can influence/promote excessive cell
division resulting in tumour growth (Hirota and Suttajit,
1988; Van Damme, 2001; Vogg et al., 1999). Also latex
of E. tirucalli and Euphorbia royleana is known to cause
conjunctivitis on contact with eyes (Shlamovitz et al.,
2009; Van Damme, 1989).
However, some members are very useful substances.
Since time immemorial, many Euphorbiaceae have been
popular traditional medicinal herbs. Genus Euphorbia and
indeed family Euphorbiaceae were named in honour of a
Greek physician to King Juba II of Mauritania called
654 J. Med. Plant. Res.
Euphorbus believed to have used Euphorbia resinifera
latex to cure ailments for example, when the King had a
swollen belly (Lovell, 1998; Van Damme, 2001). As early
as 2 BC, Euphorbiaceae such as Croton oblongifolius
Roxb. and Croton tiglium Willd. were used to cure liver
diseases, sprains, snake bites, and as a purgative for the
first as well as insanity, convulsions, asthma, tumors,
rheumatism for the latter, as documented in the Indian
Ayurveda medicine system (Kapoor, 1989). Hooper
(2002) also reports the use of Euphorbia polycarpa,
Euphorbia hirta, and Acalypha indica L. for treatment of
different ailments in the ancient Ayurveda system. In
ancient Chinese medicine, Lai et al. (2004) report 33
species belonging to 17 genera of Euphorbiaceae used in
herbal medicine. Similar reports have been cited for the
ancient Yucatan herbal system applying different
Euphorbiaceae like Euphorbia ptercineura for asthma
and cough; Croton peraeruginosus for pimples and
Phyllanthus micrandrus Müll. Arg. for wounds,
inflammations and infections among others (Ankli et al.,
1999).
Even today, many Euphorbiaceae plant concoctions,
fresh latex and teas are used in alternative medicine. For
example, E. tirucalli is known for its curative features
against diseases like warts, cancer, gonorrhea, arthritis,
asthma, cough, earache, neuralgia, rheumatism,
toothache, excrescences, tumours and others (Cataluna
and Rates, 1999; Duke, 1983; Van Damme, 1989).
Euphorbia thymifolia is used as an anti-viral against
simplex virus-2 (Gupta, 2007) whereas E. maculata is
said to cure cholera, diarrhea and dysentery
(www.botanical.com.). The latter website lists a number
of Euphorbiaceae with varying curative features
including: Euphorbia. peplus L., Euphorbia. peploides,
Euphorbia pilosa, Euphorbia palustris being remedies for
hydrophobia; Euphorbia peplus, Euphorbia helioscopia,
Euphorbia humistrata, Euphorbia hypericifolia, Euphorbia
portulacoides L., Euphorbia iata Engelm, Euphorbia
marginata Pursh, Euphorbia drummondii and Euphorbia
heterodoxa for general home ailments. Most of the
species, however, are cited in folk medicine where their
dosage and efficacy are not clear hence the need for
medical research to establish their safety.
Research has shown that some Euphorbiaceae are
actually potent as medicinal plants and their extracts
have been isolated and patented as modern drugs. A
table of some US patents is shown in Table 1.
Some of the extracts are registered drugs and as such
available on the market. Examples include Euphorbium
(resiniferatoxin), from latex of Euphorbia resinifera
(Appendino and Szallasi, 1997) marketed as ‘Complexe
Lehning Euphorbium N 88’ and used as a nasal spray or
compositum against viral infections, rhinitis of various
origins, sinusitis, chronic nasal discharge, dry and
inflamed nasal membranes as well as flu symptoms.
Euphorbia pilulifera (the asthma weed) extract has been
cited in Steadman’s drugs list and can be applied against
asthma, coryza and other respiratory infections and as an
anti-spasmodic (www.drugs.com). Dysenteral® is an
extract from Euphorbia hirta and is used in the treatment
of diarrheal diseases (Elujoba et al., 2005) while Radix is
an extract from Euphorbia kansui roots used as a
purgative. Many other Euphorbiaceae are prospective
important veterinary and agricultural biocides as
demonstrated in literature (Table 2).
Other uses of Euphorbiaceae include biodiesel
production e.g. E. tirucalli (Duke, 1983; Van Damme,
2001), Euphorbia lathyris (Duke, 1983); J. curcas (Achten
et al., 2008; de Oliveira et al., 2009; Kaushik et al., 2007;
Kumar and Sharma, 2005); M. esculenta (Adeniyi et al.,
2007); R. communis (Benavides et al., 2007; Meneghetti
et al., 2007) among others. Others are sources of food
e.g. M. esculenta (Aloys and Ming, 2006), starch e.g. M.
esculenta (Sanchez et al., 2009; Srinivas, 2007), while
others are ornamental due to their attractiveness such as
Euphorbia milli, tirucalli (Van Damme, 1989 and 2001),
Euphorbia obesa and Euphorbia pulcherrima. Other
minor uses include production of fuel wood, curving of
wooden crafts, use as hedge/fence plants, timber
production, use in re-forestation programs and others.
The above account indicates that in addition to other
uses mentioned, Euphorbiaceae is an important source
of herbal medicine of human, veterinary and agricultural
importance. The outstanding question is ‘Why is this
family significant as a medicinal taxon?’ The objective of
this review is to attempt to provide an explanation why
this may be so.
WHY IS EUPHORBIACEAE RICH IN MEDICINAL
COMPOUNDS?
Euphorbiaceae s.l. is composed of five subfamilies, 49
tribes, 317 genera and about 8,000 species (Webster,
1994). This makes it one of the biggest plant families with
probably the highest species richness in many habitats.
The implication is that in absolute terms, there is a higher
probability of having more species that are medicinal in
that one family as compared to other families. Although
such comparison (as to which family has the highest
number of medicinal species) may not have been done, it
appears that Euphorbiaceae may not compare badly to
other families. Cited inventories in different parts of the
world reveal that in Kenya, of 900 medicinal species
recorded, about 60 belong to Euphorbiaceae (Leakey,
2006), in Loja province (Southern Ecuador) they are 11
species of 214 (Bussmann and Sharon, 2006), in Jinja
district (Eastern Uganda) they are 5 out of 88 (Bukenya-
Ziraba and Kamoga, 2007), in Sango bay area (Southern
Uganda) they are 14 out of 186 (Ssegawa and
Kasenene, 2007), in Riau province, Sumatra, Indonesia,
they are 11 out of 114 (Grosvenor et al., 1995). This
approximates to about 7% of the species cited. Bearing in
mind that there are about 350 families in the plant
Mwine and Van Damme 655
Table 1. Examples of US patents of medicinal Euphorbiaceae extracts.
Patent no. Inventor Claim/ailments Species involved Patent date
US 5707631 Advanced plant Pharm. Inc. Therapeutic herbal
composition
E. lathyris January 1998
US 6844013 Peplin Biotech Pyt. Immuno-stimulation E. peplus, E. hirta,
E. drummondii
March 2001
US 2003/0165579 A1 LaRiviere Grubman and
Payne LLP. Tumour inhibition E. antiquorum February 2002
US 6432452 Peplin Biotech Pty. Anti-cancer
compound
E. peplus, E. hirta,
E. drummondii
August 2002
US 2003/0171334 A1 Peter Gordon Parsons Prostate cancer E. aaron-rossii,
E. tirucalli,
E. tomentella,
E. tomentosa
September 2003
US 6923993 Nicholas Dodato Anti-cancer
components
E. obesa August 2005
US 2007/0248694 A1 PhytoMyco Research Corp. Anti-inflammatory
properties
E. hirta October 2007
US 2006/0198905 A1 Rajesh Jain and others Ano-rectal and
colonal diseases
E. prostrata May 2008
Source: United States Patent and trademark office (seen at http://patft.uspto.gov.)
Table 2. Examples of pesticidal species in Euphorbiaceae and their remedies.
Pesticidal
feature
Species Chemical compound(s) Cited reference(s)
Anti- bacterial
E. guyoniana Boiss. and Reut. Diterpenes El-Bassuony (2007)
E. sororia Schrenk Ceramides and ellagic acid derivatives Zhang et al. (2008)
E. hirta Tannins, alkaloids and flavonoids Ogbulie et al. (2007)
E. pubescens Vahl. Diterpenes and ent-abietanes Valente et al. (2004)
M. esculenta Glycocide Zakaria et al. (2006)
E. sessiliflora Roxb. Triterpenes and ellagic acid derivatives Sutthivaiyakit et al. (2000)
E. segetalis L. Coumarins and steroids Madureiira et al. (2002)
J. podagrica Hook. Diterpenoids Alyelaagbe et al. (2007)
E. ebracteolata Hayata Casbane diterpenoids Xu et al. (1998)
E. heterophylla Saponins, flavonoids and tannins Falodun et al. (2008)
Drypetes inaequalis Hutch. Triterpenoid esters and saponins Awanchiri et al. (2009)
Anti-viral
E. kansui Triterpenes, sterols and diterpenes Zheng et al. (1998)
E. hyberna L. Diterpenes Bedoya et al. (2009)
E. cotinifolia L. , E. tirucalli Diterpenes Bentacur-Galvis et al. (2002)
E. thymifolia Alkaloids Jabbar and Khan (1965)
E. thymifolia Triterpenes and alkaloids Lin et al. (2002)
656 J. Med. Plant. Res.
Table 2. Contd.
Anti-fungal
Macaranga monandra Müll.Arg. Diterpenes Salah et al. (2003)
E. nivulia Buch.-Ham. - Annapurna et al. (2004)
E. hirta, E. tirucalli Diterpenes and triterpenes Mohamed et al. (1996)
R. communis Fatty acids Maria Fatima et al. (2004)
J. curcas Glucanase protein Wei et al. (2005)
E. tirucalli, E. helioscopia, E.
splendens Bojer. Ex Hooke. E.
pulcherrima Willd. Ex Klotzsch.
Diterpenes Devi and Gupta (2000)
E. pulcherrima - Cox et al. (2006)
Nematicidal Phyllanthus niruri L. Flavanones Shakil et al. (2006); Shakil et
al. (2008)
E. kansui Diterpenes and ingenane Shi et al. (2007); Shi et al.
(2008)
E. hirta Phenols Adedapo et al. (2005)
J. podagrica Peptides Dahiya (2008)
Moluscicidal
E. tirucalli - Jurberg et al. (1985);
Vassiliades (1984)
E. conspicua N.E. Br. Diterpenes and triterpenes Dos Santos et al. (2007)
E. splendens - de Vasconcellos and de
Amorim (2003)
J. elliptica Müll. Arg. Diterpenes dos Santos and Sant’Ana
(1999)
J. curcas Phorbal esters Gubitz et al. (1999)
E. paralias L. Diterpenes Abdelgaleil et al. (2002)
Insecticidal
E. hirta Flavonol glycosides Liu et al. (2007)
J. curcas Sterols, triterpenes alcohols and
acids Adebowale and Adedire
(2006)
R. communis Flavonoids Shripad (2003)
R. communis Ricinine Maria Fatima et al. (2004)
J. curcas Diterpenoids Goel et al. (2007)
C. pseudoniveus, C. suberosus Essential oils Perez-Amador et al. (2003)
Anti-
leishmanial
D. chevalieri Furansesquiterpene and
triterpenoids Wansi et al. (2007)
J. grossidentata, J. isabellii diterpenes Schmeda-Hirschmann et al.
(1996)
J. grossidentata Diterpenes Akendengue et al. (1999)
P. cajucara Essential oils Ahmed et al. (2006)
Kingdom, this is not a bad score. This, however, depends
upon the region in question since Euphorbiaceae is most
prevalent in tropical and subtropical areas.
According to Oldfield (1997), a good number of
Euphorbiaceae species especially of genus Euphorbia
(650 species), are succulent. The latter author describes
succulence as a plant characteristic mainly tropical or
subtropical that has to conserve water due to habitat
aridity. Von Willert et al. (1990) describe it as a
characteristic that makes a plant temporarily independent
from external water supply when soil water conditions
have deteriorated such that the roots are no longer able
to provide the necessary water from the soil. They argue
that this is only a temporary adaptation to aridity and
unless conditions allow the refilling of the plant’s
succulent tissues, it will not survive. This implies that
such plants tend to avoid mechanisms that result into
water loss (including thin leaf surface, broad leaves and a
large number of leaves), whereas they invest in features
that conserve water such as thick waxy leaves, scaly
leaves or thorns, fewer/sunken stomata, use of stems for
photosynthesis, use of Crassulacean Acid Metabolism
(CAM) and others. It is therefore a survival strategy for
plants in arid and semi-arid areas.
According to Griffiths et al. (2008), leaf succulence is a
key morphological correlate of the capacity for CAM,
since succulence increases the plants’ commitment to the
use of CAM pathway during carbon dioxide fixation. In
their experiments with two succulent plants, the latter
authors found out that the magnitude of CAM was higher
for the more succulent leaves of Kalanchoe
daigremontiana Raym. Hamet and H. Perrier
(Crassulaceae) compared to the less succulent leaves of
Kalanchoe pinnata (Lam.) Pers. In the same spirit, Van
Damme (1989) had earlier on pointed to the CAM-
succulence syndrome stating that the two have to always
go together, to which Lüttge (2004) concurs insisting that
all CAM plants display some level of succulence.
The CAM pathway is known to be under circadian
control and is subject to regulation by multiple oscillators,
which modulate elements of the pathway in line with
environmental conditions (Borland et al., 1999; Borland
and Taybi, 2004). In line with this, Lüttge (2004 and
2008) enumerates a ‘wealth of environmental factors’
known to determine, or at least modulate the expression
of CAM to include: carbon dioxide, water, absolute
temperatures, day-night temperature regimes, irradiance
and salinity among others.
The latter author goes on to provide a model that
relates these factors, drawing a conclusion that CAM-
prone ecosystems are those that are governed by a
network of interacting stress factors requiring versatile
responses and not systems where a single stress factor
strongly prevails. They point out a number of CAM
domains or ecosystems that are likely to host
CAM/succulent plants including submerged aquatic sites,
deserts, salinas, savannahs, inselbergs, forests, and high
latitudes such as tropical highlands and alpine regions.
This implies that although the CAM pathway is an
adaptation for succulent plants to balancing their carbon
and water budgets (Lüttge, 2008; Von Willert et al.,
1990), it is also a survival mechanism for adverse
conditions. Indeed, the extent of succulence has been
positively correlated to both colonization of increasingly
arid habitats and an increased contribution of CAM
activity to total carbon gain (Herrera, 2009; Kluge et al.,
2001, Van Damme, 1989 and 2001), implying that
succulent plants are better equipped for new habitat
colonization than their non- or less succulent relatives,
which gives them higher chances for survival in a wider
habitat range.
Mwine and Van Damme 657
As an extra adaptation, some succulent plants have been
found to combine both C3 and CAM pathways. For
example, Van Damme (1989 and 2001) reports that E.
tirucalli utilizes both pathways. Its minute deciduous
leaves utilize the C3 pathway while the stem uses CAM.
He explains that the small leaves are preferentially used
in normal situations whereas the green stem takes over
when the leaves fall off in arid conditions. According to
him, a combination of both pathways increases the
plant’s water use efficiency (WUE) since the small leaves
have a high affinity for carbon dioxide but tend to use
water less efficiently. He predicts that there could be
many other Euphorbiaceae using the same mechanism
which he says improves productivity and ability to
colonize a wide range of habitats.
Euphorbiaceae are very widely distributed in almost all
habitats and occupy a wide range of climatic and soil
disparities. Ahmad et al. (2006) and Bloomquist (2004)
report that different habitat conditions e.g. soils, pH,
temperatures and moisture tend to influence plant
physiological processes hence the manufacture and
accumulation of different chemical substances. In
agreement, Melten et al. (2009) confirm that plants’
responses can differ due to different factors. For
example, physiological alteration of photosynthetic
enzyme ratios to adjust to changing light conditions,
suberin production to limit moisture loss from roots,
genetic regulation of enzymes in response to resource
limitation, or production of secondary metabolites in
leaves in response to insect/microbial attack. In the same
view, Veronese et al. (2003) found out that different
herbivores (as found in different environments) tend to
induce action of different defense systems. The
implication of all this is that due to a wide range of
conditions, which different Euphorbiaceae species are
subjected to, the latter tend to manufacture a wide range
of secondary plant substances to aid response to a
disparity of stimuli in their particular habitats. For
example, Zhang et al. (2000) reported manufacture of
different lectins due to different environmental stress
factors that occur in varying habitats. Lectins constitute
part of a plant’s defense system against herbivores (Van
Damme, 2008). Similarly, Agrawal and Rutter (1998)
found that changes in environmental cues can trigger
modifications in a plant’s defense strategy as was
witnessed in ant plants (Myrmecodia spp.). Also,
Taniguchi et al. (2002) established that production of
secondary substances like pentagalloylglucose was
remarkably enhanced under light irradiation compared to
dark conditions while tannin production was greatly
affected by changing the concentrations and composition
of nitrogen sources. These and other findings support the
view that Euphorbiaceae may have a variety of medicinal
substances due to a disparity of environmental conditions
(stress factors) accruing from a wide habitat range.
Biosynthesis of secondary metabolites is a complex
process and is still poorly understood. Hadacek (2002)
658 J. Med. Plant. Res.
states that biosynthesis and accumulation of secondary
metabolites arise from highly regulated processes
requiring both genetic and environment-specific controls.
In a related view, Cavalier-Smith (2007) says that
secondary metabolites are produced from universally
present precursors mostly acetyl-CoA, amino acids or
shikimate (shikimic acid) by taxon specific enzymes – the
reason why most secondary metabolites are restricted to
a single major taxon on the universal phyllogenetic tree
or evolutionally related taxa. This specificity has been
shown and supported by modern molecular techniques,
for example, using cytochrome P450 enzymes (CPY79A1
and CYP71E1) involved in biosynthesis of the cynogenic
glucoside dhurrin in Sorghum bicolor L. (Kahn et al.,
1999). This phenomenon is not new in the plant kingdom
because some secondary metabolites are considered to
be so taxon specific as to be used in classification of
certain taxa or acting as proof of monophylly
(Gershenzon and Mabry, 1983; Herbert, 1989; Seigler,
1994; Wink, 2003). What may be controversial, however,
is that there is a growing volume of evidence to show that
plants of the same genus or family may synthesize
different or at least varying secondary metabolites when
growing in different conditions (Figueiredo et al., 2008;
Koricheva et al., 1998; Wink, 2003). Such inconsistencies
in secondary metabolite profiles have been attributed to
among other factors, differential expression of
corresponding genes (Wink, 2003) but also change in
gene sequence and mutations (Theis and Lerdau, 2003).
These genetically related factors act in the wake of
varying environmental factors, to cause alterations in
secondary metabolite assemblages reflecting adaptations
and particular life strategies embedded in a given
phylogenetic framework (Koricheva et al., 1998). As
indicated by Ogunwenmo et al. (2007), this argument
appears plausible for the case of widely distributed
families like Euphorbiaceae. Varying secondary metabo-
lites may be synthesized within the taxon, as a result of
different gene expression and increasing mutation loads,
accruing from stressful environments that characterize
most Euphorbiaceae habitats. This may result into a
richer assemblage of secondary metabolites within the
family.
During their evolution, CAM plants have developed a
number of anatomical, physiological and genetic
changes/adaptations, which differentiate them from C3
plants. Cushman and Bohnert (1997) mention several of
them such as thin-walled cells containing prominent
vacuoles, varying degrees of succulence and capacity to
handle high degrees of organic acid accumulation. The
latter authors go on to say that just like evolution
processes, CAM induction or process of shifting from C3
to CAM, involves the regulation of a variety of enzymes
and metabolite transporters making it a very complex
metabolic adaptation (to environmental stress). They give
an example of increased activities of glycolytic,
gluconeogenic and C4 acid metabolism enzymes
including phosphoenolpyruvate carboxylase (PEPC),
phosphoenolpyruvate carboxykinase (PEPCK) and
pyruvate phosphate dikinase (PPDK) which increase 40
fold due to environmental stress. On the genetic side, the
same authors, citing transcription essays with nuclei
isolated from leaves of Mesembryanthemum crystallinum
L. (Aizoaceae), show that CAM-specific genes increase
two to six times when plants are exposed to high salinity
(a stress factor). In support, Borland and Taybi (2004),
note that although physiological processes associated
with this high organic acid accumulation are energy-
intensive, the potential for high productivity is not
compromised. To substantiate this fact, they give
examples of agronomically important CAM species
including pineapple (Ananas comosus Mill.) and Agave
spp. that show productivities rivaling that of C3 and C4
plants. Since organic acids and genes are responsible for
physiological plant processes, this implies that CAM
plants are likely to have higher productivity both in
quantity and quality (variety) of chemical substances
including enzymes, proteins, amino acid as well as
secondary metabolites of various nature.
Possession of a variety of chemical substances may
entail being rich in medicinal attributes. For example,
over sixty jatrophane, modified jatrophane, segetane,
pepluane and paraliane diterpenoids were extracted,
purified and characterized from different Euphorbiaceae
such as Euphorbia dendroides, Euphorbia characias,
Euphorbia peplus, Euphorbia amygdaloides, and
Euphorbia paralias. Based on jatrophane and modified
jatrophane skeletons these were shown to be potent
inhibitors of P-glycoprotein activity – a membrane protein
that confers upon cells the ability to resist lethal doses of
certain cytotoxic drugs by pumping them out of the cells,
thus reducing cytotoxic effects (Barile et al., 2008).
Similarly, Corea et al. (2005) report the discovery of two
new diterpenes, pepluanone 1 and 2 from E. peplus,
which act as anti-inflammatory agents. Also Falodun et
al. (2008) identified secondary metabolites such as
saponins, flavonoids and tannins from E. heterophylla
which exhibited good activity against xanthine oxidase
enzymes. Related findings are distributed in the whole
Euphorbiaceae literature (Table 3).
CONCLUSION
Each plant family may have its own good reason for
possession of medicinal properties. For Euphorbiaceae
family members, it would appear that their diverse
medicinal properties are associated with their wide
distribution which is supported by their survival
adaptations such as succulence and CAM pathway. The
exposure to a wide range of habitats predisposes them to
inevitably high mutation loads (accruing from stressful
habitats) and a large range of environmental stimuli
hence the necessity to develop a wide battery range of
defensive secondary metabolites. These issues may
explain why the family is widely pharmaceutical. These
Mwine and Van Damme 659
Table 3. Chemical substances found in Euphorbiaceae and their pharmaceutical indications.
Chemical substance Medicinal indication Cited reference(s)
Diterpenes
Anti-tumor Duarte et al. (2008); Konoshima et al. (2001); Krebs et al. (2004)
Anti-biotic El-Bassuony (2007); Li et al. (2008); Mathabe et al. (2008)
Anti-fungal Salah et al. (2003)
Anti-plasmodial Attioua et al. (2007)
Anti-ulcergenic Hiruma-Lima et al. (2002)
Trypanacidal Schmeda-Hirschmann et al. (1996)
Triterpenes
Anti-biotic Awanchiri et al. (2009); Mathabe et al. (2008)
Vaso-depressor Barla et al. (2006)
Anti-inflammatory Canelon et al. (2008); Nkeh et al. (2008)
Analgesic Nkeh et al. (2003)
Anti-fungi Ekpo and Pretorius (2007)
Flavonoids Anti-malarial Liu et al. (2007)
Anti-inflammatory Ekpo and Pretorius (2007)
Saponins Cytotoxic Kiem et al. (2009)
Anti-ulcer Ukwe (1997)
Tannins
Anti-septic Ekpo and Pretorius (2007)
Anti-viral Bessong et al. (2006); Liu et al. (1999)
Anti-mutagenic Rossi et al. (2003)
Anti-fungal Hwang et al. (2001)
Alkaloids Anti-microbial Dias et al. (2007); Gressler et al. (2008)
Anti-tumor Suarez et al. (2004)
Esters
Anti-tumor Blanco-Molina et al. (2001); Goel et al. (2007)
Anti-biotic Goel et al. (2007)
Cytotoxic Baloch et al. (2006)
Allergic reactions Thumm et al. (2002)
Cancerous Cataluña and Rates (1999)
Ricin
Cytotoxic Lombard et al. (2001)
Lipolytic Lombard et al. (2001)
Invertase activation Vattuone et al. (1991)
Phenols Anti-tumor Yu et al. (2005)
Anti-oxidant Yang et al. (2007)
literature review findings compare well with reports of
other workers like Ahmad et al. (2006) and Okgibo et
al. (2009) who similarly but independently reviewed a
variety of Euphorbiaceae-based phytochemi-cals
including alkaloids, phenols, flavonoids, saponins,
tannins and essential oils and described their origins,
characteristics and therapeutic uses.
This review has revealed a rich variety of medicinal and
potentially medicinal properties of Euphorbiaceae and
attempted to elucidate why Euphorbiaceae tick as
medicinal plants. Euphorbiaceae is therefore, a good
starting point for a search for phytomedicines of human,
veterinary or pesticidal nature.
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