Nutritional and pharmacological potential of the genus Ceratotheca—
An underutilized leafy vegetable of Africa
Nqobile A. Masondo, Jeffrey F. Finnie, Johannes Van Staden
Research Centre for Plant Growth and Development, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg Campus, P/Bag X01, Scottsville
3209, South Africa
Received 25 August 2015
Received in revised form
30 November 2015
Accepted 1 December 2015
Available online 4 December 2015
Traditional leafy vegetables
Ethnopharmacological relevance: Ceratotheca (Pedaliaceae) is an endemic African genus comprising of
ﬁve species. The genus is commonly used as a leafy vegetable with medicinal properties.
Aim of the review: The review aims to highlight the unexplored nutritional and pharmacological po-
tential of African indigenous leafy vegetables Ceratotheca sesamoides and triloba, in order to conserve and
domesticate these species.
Methods: The information was obtained from various search engines such as Scopus, Google Scholar and
Web of Science as well as Ethnobotanical books.
Results: Ceratotheca sesamoides and triloba have good nutritional potential. The species are high in en-
ergy levels, fat content, proteins and carbohydrate values. The species have also been reported to have
good antibacterial, antidiarrhoeal, antidiabetic, antiplasmodial and antiviral properties. These species
have slight toxicity and cytotoxic activity when extracted at high concentrations but no mutagenic ac-
tivity was detected.
Conclusions: To date, few studies have documented the usage (nutrition and pharmacology) of Cer-
atotheca sesamoides and triloba. More studies investigating the nutritional content as well as methods of
its improvement are necessary if the plant is to be included as a domesticated vegetable crop. Safety and
toxicity analysis of this leafy vegetable need to be extensively studied as the plants are consumed in high
&2015 Elsevier Ireland Ltd. All rights reserved.
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
2. Botanical distribution and general morphology of Ceratotheca species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
2.1. Ceratotheca sesamoides ......................................................................................... 211
2.2. Ceratotheca triloba ............................................................................................. 211
3. Consumption of Ceratotheca species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
4. Nutritional value of Ceratotheca species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
5. Medicinal properties of Ceratotheca species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
6. Phytochemistry of Ceratotheca species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
7. Pharmacological properties of Ceratotheca species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
7.1. Antioxidant activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
7.2. Antimicrobial and antiviral activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
7.3. Antidiarrhoeal activity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
7.4. Anti-inﬂammatory activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
7.5. Anti-diabetic activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
7.6. Antiplasmodial activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
7.7. Hyaluronidase, phospholipase and proteolytic activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
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0378-8741/&2015 Elsevier Ireland Ltd. All rights reserved.
E-mail address: email@example.com (J. Van Staden).
Journal of Ethnopharmacology 178 (2016) 209–221
7.8. Other documented properties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
8. Safety and toxicity of Ceratotheca species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
9. Propagation strategies of Ceratotheca species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
10. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
The Pedaliaceae is primarily an African family which comprises
of 11 genera. The family is characterised by a distinct type of
mucilage hair, covering most parts of the plant including the
leaves. The mucilage glands secretes mucilage once in contact with
water (Ihlenfeldt, 2004). Amongst the genera is Ceratotheca, which
consists of ﬁve species endemic to Africa. The generic name Cer-
atotheca was derived from the Greek names kerato (horned) and
theke (a case) describing the plants discrete sharp horned capsule
(Duncan, 2011). The genus has been documented to closely re-
semble Sesamum L. which produces sesame seeds and oil, with
minor differences (Bedigian, 2004).
Among the ﬁve species, Ceratotheca sesamoides Endl. and Cer-
atotheca triloba (Bernh.) Hook.f. are the only two species docu-
mented for their nutritional and pharmaceutical importance. Even
though the species have an acrid smell and hairs on their leaves,
they are still included in the diet (commonly classiﬁed as tradi-
tional leafy vegetables) in some parts of Africa. C. sesamoides has
been reported to be a good source of proteins, sugars, and calcium
(Fasakin, 2004;Grubben and Denton, 2004;Yadang et al., 2009),
whilst C. triloba contains high energy (kJ), lipids and magnesium
levels (Odhav et al., 2007). Apart from their nutritional role in
many diets, the two species are also widely utilized in traditional
medicine in the treatment of various ailments. For instance, in
parts of Nigeria, C. sesamoides is used in the treatment of diar-
rhoea, conjunctivitis, snakebites, skin diseases and facilitatation of
birth in both humans and animals (Grubben and Denton, 2004). In
South Africa, C. triloba leaves are extensively utilized by the Zulu
people for the relief of painful menstruation, stomach cramps and
diarrhoea (Pooley, 1998;Watt and Breyer-Brandwijk, 1962). Con-
sequently, Ceratotheca species have been evaluated in both in vitro
and in vivo studies for antimicrobial, antidiarrhoeal, anti-in-
ﬂammatory and anti-diabetic properties (Akula and Odhav, 2008;
Mohanlall and Odhav, 2013;Odhav et al., 2010;Toyin et al., 2012).
Traditional vegetables such as C. sesamoides and C. triloba are often
classiﬁed as functional foods due to their health beneﬁts extending
beyond basic nutritional value (Van Wyk and Gericke, 2000). Al-
though the importance of these plants in Africa remains in-
adequate and poorly documented, the available literature supports
their high and valuable nutritional status as well as their phar-
macological value. Thus, this review aims to unravel the un-
explored potential and signiﬁcance of the African endemic species
C. sesamoides and C. triloba, and their nutritional and pharmaco-
Fig. 1. Ceratotheca morphology: A—whole plant; B—ﬂowers; and C—seed capsule.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221210
2. Botanical distribution and general morphology of Cer-
The ﬁve species of Ceratotheca include the annual Ceratotheca
integribracteata Engl. distributed in Angola and Namibia; the an-
nual Ceratotheca saxicola E.A.Bruce endemic to Soutpansberg
Centre, Limpopo Province, South Africa; the annual C. sesamoides
Endl. endemic to the northern parts of West Africa; the annual and
biennial C. triloba (Bernh.) Hook.f. which is distributed in southern
Africa and the perennial Ceratotheca reniformis Abels found in
The genus consists of erect plants which grows to a height of
0.6–2.0 m. Ceratotheca leaves are opposite with lower leaves larger
and the uppermost leaves narrowly to broadly lanceolate. The
ﬂowers have a foxglove-like shape, produced in pairs varying from
white, pink or mauve (Fig. 1). Flowering season commences
around late January (summer) until May (autumn), depending on
the species. The plant produces distinctive horn shaped seeds that
are dark brown or black in colour (Duncan, 2011). Similar to other
Pedaliaceae members, Ceratotheca species are covered with mu-
cilage glands. These glands allow the plants to withstand severe
dehydration without tissue damage, making them drought re-
sistant (Bedigian and Adetula, 2004).
2.1. Ceratotheca sesamoides
C. sesamoides commonly known as ‘false sesame’was ﬁrst de-
scribed by Prof. Stephan Endlicher (1804–1849), Director of the
Botanical Garden of Vienna in Linnaea (Grubben and Denton,
2004). The species has two synonyms, Ceratotheca melanosperma
Hochst. ex Bernh. (1842) and Sesamum heudelotii Stapf (1906). The
plant is also known by various names such as Eku (Yoruba-Wes-
tern Nigeria), Bungu (Nigeria), Tchaba-laba (Guinea Bissau) and
Lalu-caminho (Senegal). The species is found in West Africa and
has extended its distribution from Senegal to Tanzania, Democratic
Republic of Congo, southwards to Botswana, Mozambique, Zim-
babwe and Zambia. Even though the species grows as a weed, it
has become one of the most frequently cultivated leafy vegetables
in most parts of sub-Saharan Africa. The plant is easily adapted to a
wide range of environments including open grasslands and tree
savanna with well-drained sandy soils (Grubben and Denton,
2004). It grows well in sunny habitats making it heat and drought
2.2. Ceratotheca triloba
C. triloba commonly known as African foxglove was ﬁrst
documented in southern Botswana (1812) by William Burchell at
Chue Spring (now Heuningvlei). It was initially recorded as Spor-
ledera triloba by the German botanist Johann Bernhardi (1774–
1850), then described as Ceratotheca by J.D. Hooker in Curtis’s
Botanical Magazine (Duncan, 2011). Its synonyms include Cer-
atotheca lamiifolia (Engl.) Engl., Sesamum lamiifolium Engl., Spor-
ledera kraussiana Bernh., and S. triloba Bernh. (Raimondo et al.,
2009). The species is commonly referred to as Wild foxglove
(English); Vingerhoedblombossie (Afrikaans); Udoncalwabathwa,
Udonqabathwa (Zulu); Mudyangaringa, Nyamanhuwe (Shona). C.
triloba is widely distributed in southern Africa namely South
Africa, Mozambique, Zimbabwe, Zambia and Angola (Pickering and
Roe, 2009). The plant either grows as a weed or is cultivated in
some regions. In South Africa the plants grow best in summer
rainfall regions such as grasslands. Seeds germinate in spring or
early summer, in rich and well-drained soil and ﬂowers before the
onset of winter.
3. Consumption of Ceratotheca species
In the last two decades, there has been a growing interest in
the potential of traditional vegetables in overcoming the nutri-
tional gap experienced by the population in developing countries.
However, several leafy vegetable species are yet to be recognized
and documented for the important role they play in human health.
In addition, data collection on leafy vegetables has been poorly
characterized within international genebanks (Maggioni, 2004).
Due to such factors, knowledge on the usage (nutritional and
medicinal) of neglected or underutilised crop species is crucial for
C. sesamoides is amongst the important African indigenous
vegetables widely utilized especially for the oil it produces. Al-
though the plant has been reported to be widely consumed in
most parts of West Africa (Bedigian and Adetula, 2004), available
literature on its consumption is still insufﬁcient (Table 1). Fur-
thermore, the available literature has described the consumption
of C. sesamoides as being dependent on its distribution and scarcity
of other preferred food sources in the region. For instance, Mertz
et al. (2001) reported on the plant as being the least consumed
leafy vegetable in Burkina Faso even though it has a pleasant taste.
Dansi et al. (2012) also found that the species was abundant in
Benin yet its consumption was insigniﬁcant compared to other
leafy vegetables. Even when cultivated, C. sesamoides plants were
reported to have a low consumption rate in some regions (Lykke
et al., 2002;N’Danikou et al., 2011). Nevertheless, there are some
regions that have documented the importance of C. sesamoides in
the diet. In Benin (Sudano-Guinean and Sudanian region), the
plant is amongst the most commonly consumed non-cultivated
leafy vegetables (Achigan-Dako et al., 2011). The species has also
been reported as one of the most frequently consumed vegetable
with millet in south-western Niger (Ayantunde et al., 2009). The
plant also serves as feed for livestock. The species has been re-
corded to be a signiﬁcant wild crop in Zambia, with 37% of
households reporting its consumption (Bedigian, 2004). Unlike C.
sesamoides, literature on the consumption of C. triloba in southern
Africa is sparse. The species has thus far been recorded to be
consumed in drier areas of Zimbabwe and the leafy vegetable was
classiﬁed as a ‘poor man’s food’(Maroyi, 2011). The infrequent
consumption of C. sesamoides and C. triloba in some areas might be
due to several reasons such as preference for other vegetables,
availability/seasonality of the species and the leafy vegetable being
considered as “poor man’s food”. Nonetheless, these vegetables
may serve to supplement the mainly starch-based diets of many
under-developed communities in Africa since they can grow in the
wild and are also easily cultivated.
4. Nutritional value of Ceratotheca species
Nutrient-rich foods which are a good source of proteins, vita-
mins, minerals, ﬁbres and amino acids are vital for growth and
development in children and adults. The recommended dietary
allowance (RDA) for minerals in adults is as follows: calcium,
1000 mg/day; phosphorous, 800 mg/day; copper, 900 mg/day;
zinc, 10 mg/day; magnesium, 400 mg/day; manganese, 7 mg/day;
and iron, 8 mg/day (Odhav et al., 2007). Although the importance
of nutrients such as proteins is well-recognized, most of the po-
pulation in developing countries still lack the daily required
amount in their diets. Hence, several studies have reported on the
value of vegetables as major constituents of micro/macronutrient
as well as dietary energy (Aletor et al., 2002;Barminas et al., 1998;
Uusiku et al., 2010). Thus, supplementing poor diets with vege-
tables can contribute greatly to the reduction of malnutrition in
the majority of the population in sub-Saharan Africa (Bedigian,
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221 211
2004;Van Wyk and Gericke, 2000).
During the preparation of C. sesamoides and C. triloba, most
parts of the plant are consumed including the leaves, ﬂowers and
seeds. C. sesamoides is prepared by grinding leaves into ﬁne
powders and mixing them with groundnut ﬂour, salt and hot
water then cooking for a few minutes. The mixture is eaten as a
sauce with porridge. Onion and tomatoes can be added in order to
soften and reduce the bitterness of the leaves (Grubben and
Denton, 2004). Crushed seeds of C. sesamoides can be eaten with
beans or cassava and the oil added in salads (Grubben and Denton,
2004). The leaves of C. sesamoides are frequently served with other
leafy vegetables, possibly due to their laxative effect (Bedigian,
2004). C. triloba is cooked as a spinach. The unpleasant smell of
leaves disappears once boiled. The vegetable has a sweet taste and
is also included as a relish in some dishes (Tredgold, 1986).
The nutritional composition of C. sesamoides and C. triloba is
presented in Table 2. Leaf extracts of C. sesamoides were reported
to be a good source of proteins, carbohydrates, minerals and en-
ergy (kJ) (Bedigian and Adetula, 2004). In addition, the seeds of the
plant demonstrated even better/higher energy, protein, fat, car-
bohydrate and calcium contents compared to the leaf extracts. The
seeds of C. sesamoides have been recorded to yield 35% oils when
extracted with petroleum ether (Bolton, 1919). The author found
that upon extraction, C. sesamoides seeds tested negative against
Baudouin reaction colour test similar to Sesamum indicum seeds. In
addition, results from other tests (e.g. free fatty acids) showed
similarities in oil seed constituents with a slightly lower speciﬁc
gravity in C. sesamoides. The potentially good nutritional content of
either wild or cultivated C. sesamoides (leaves and seeds) has been
further validated by several studies (Fasakin, 2004;Fasola and
Ogunsola, 2014;Mitchikpe et al., 2008;Yadang et al., 2009). The
species has been reported to have a high mineral concentration as
well as high levels of provitamin-A [3681 retinol equivalents (RE)/
IOO g] (Bedigian, 2004). However, no studies have evaluated the
nutritional content of C. sesamoides ﬂowers even though they are
included in some dishes. Nutritional composition of C. triloba has
been established (Odhav et al., 2007). The species contain high
energy levels (kJ) as well as fat content, however its mineral
content (calcium, phosphorus, sodium, manganese, copper, zinc
and iron) were recorded to be very low when compared to com-
monly consumed leafy vegetables (Odhav et al., 2007).
Trypsin inhibitors are compounds that interfere with protein
digestion causing pancreatic enlargement and enhancement of
chemically induced pancreatic tumours (Venter and Van Eyssen,
2001). A study by Vanderjagt et al. (2000) evaluated the anti-
nutrient properties (compounds that interfere with the absorption
of nutrients) in C. sesamoides and reported its poor trypsin in-
hibitory activity (0.21–0.45 mg/mg dry weight). Leaves of C. sesa-
moides contained minor traces of trypsin-neutralizing activity
when compared to other leafy vegetables and commercial species
from Niger [e.g spinach (Amaranthus spinosus) 3.45 mg/mg dry
weight]. The trypsin activity observed in the leaf extracts showed
resistance to boiling with 100% heat resistance. The ability of
plants to resist change in trypsin activity is of interest since the co-
presence of protease inhibitors in ingested plants might reach the
small intestine and block the trypsin activity which catalyses the
hydrolysis of dietary proteins. This is an important process for
protein digestion and absorption especially for C. sesamoides
which has been reported to be a poor source of protein (Yadang
et al., 2009). C. triloba leaf extracts were recorded at an estimated
913 (TAU) trypsin inhibition and small traces of phytic acid
(0.04 mg/ml). The presence of phytic acid in vegetables leads to
iron inhibition in both infants and adults (Hurrell, 2003). It has
been estimated that removal of phytic acid tor0.01 mg/g en-
hances iron absorption (4–5 fold) in comparison to its addition
which results in decreased iron absorption (Hurrell et al., 1992).
Documented surveys on the consumption of Ceratotheca sesamoides and Ceratotheca triloba in African countries.
Scientiﬁc name Distribution Status of distribution Result highlights References
Ceratotheca sesamoides Endl. Benin (Sudano–Guinean and Suda-
Wild Among the 245 species surveyed in Benin, Ceratotheca sesamoides was the most frequently consumed
non-cultivated vegetable in the two regions.
Achigan-Dako et al.
Ceratotheca sesamoides Endl. Benin Wild The species is recognized as one of the most widely distributed and uncultivated leafy vegetable
indigenous to the country.
Adéoti et al. (2009)
Ceratotheca sesamoides Endl. Burkina Faso Wild Even though Ceratotheca sesamoides is the least consumed vegetable in Silmiogou and Ningaré re-
gions, it is still considered the tastiest vegetable appearing in few meals.
Mertz et al. (2001)
Ceratotheca sesamoides Endl. South-western Niger Wild The species was reported as one the four most recognized plants consumed with solid millet. The
plant was also found to be consumed by livestock.
Ayantunde et al. (2009)
Ceratotheca sesamoides Endl. Benin Wild In some parts of Benin, the availability of Ceratotheca sesamoides is abundant, however its con-
sumption is negligible.
Dansi et al. (2012)
Ceratotheca sesamoides Endl. North of Benin Cultivated The species was ranked at number 13 in order of its importance in the markets traded. Dansi et al. (2008)
Ceratotheca sesamoides Endl. Burkina Faso Cultivated Consumption of Ceratotheca sesamoides accounted for less than 1% of the meals in Silmiogou
Lykke et al. (2002)
Ceratotheca sesamoides Endl. Southern Benin (Dan forest) Wild The species occurs in different habitats and its value has been ranked at number 27 in the southern
region of Benin.
N’Danikou et al. (2011)
Ceratotheca triloba (Bernh.)
Zimbabwe Wild In the 32 edible traditional vegetables identiﬁed in Zimbabwe, Ceratotheca triloba was found to be
consumed in Beitbridge, Binga, Tanganda Halt and Umguza Districts. It has been classiﬁed as a “poor
man’s food”therefore not traded in the markets.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221212
Therefore, presence of phytic acid in such minor quantities will
allow for absorption of important minerals such as iron in humans.
5. Medicinal properties of Ceratotheca species
Globally, approximately 80% of the population continues to
utilize traditional medicine for several ailments. Thus, extensive
research focusing on scientiﬁc evaluation of traditional medicine
from plant origin has been on-going for the past few decades.
Apart from their nutritional value, leafy vegetables have long been
known to possess medicinal properties (Tredgold, 1986;Van Wyk
and Gericke, 2000;Watt and Breyer-Brandwijk, 1962). In this case,
C. sesamoides and C. triloba have been widely documented for their
utilization in traditional medicine compared to their consumption
In Nigeria, C. sesamoides is traditionally used in the treatment of
diarrhoea. Leaves are soaked in water and drops of the slimy re-
sidue are used to cure conjunctivitis. The mucilage is occasionally
used as an emollient and lubricant (Grubben and Denton, 2004). A
mixture of warmed, crushed leaves with ash aids in relieving
pressure on inﬂamed cervical lymph nodes. Infusion of C. sesa-
moides leaves with rhizomes of Anchomanes difformis (Blume)
Engl. can be applied topically for the treatment of leprosy. The
plant is used as an aphrodisiac, and in the treatment of jaundice,
snake bites and skin ailments. C. sesamoides leaf infusions are used
to facilitate delivery in both humans and animals (Grubben and
Denton, 2004). In northern Nigeria, C. sesamoides seeds are used to
relieve circumcision pains (Abubakar et al., 2007). Leaf decoctions
of C. sesamoides can either be administrated orally or in a form of a
bath for the treatment of malaria in Mali (Diarra et al., 2015). In
Baskoure, Kourittenga Province, Burkina Faso, the whole plant is
used as a decoction to cure stomach ache (Nadembega et al., 2011).
In South Africa, C. triloba is traditionally used for the relief of
painful menstruation, stomach cramps, nausea, fever and diar-
rhoea (Pooley, 1998). In Zimbabwe, the leaves of the species are
administered to induce abortion (Hutchings et al., 1996). Leaf in-
fusions of the plant aids in the relief of gastro-intestinal cramps
and ﬂatulence (Roberts, 1990;Watt and Breyer-Brandwijk, 1962).
C. triloba leaves are used as an abortifacient and for
dysmenorrhoea (Van Wyk and Gericke, 2000;Watt and Breyer-
Brandwijk, 1962). Root infusions are used for the treatment of sore
eyes and ears (Hutchings et al., 1996). When soaked, the whole
plant can be used as soap or shampoo. Due to the unpleasant
odour the plant produces, its leaves have been utilized in insect-
repellent sprays (Roberts, 1990).
6. Phytochemistry of Ceratotheca species
The phytochemicals reported from C. sesamoides and C. triloba
are summarised in Table 3. Phytochemical quantiﬁcation of C. se-
samoides leaves yielded ﬂavonoids, saponins, alkaloids, tannins
and phenols (Fasola and Ogunsola, 2014;Toyin et al., 2012). Ac-
cording to Fasola and Ogunsola (2014), different stages of growth
had a signiﬁcant impact on the phytochemical content of C. sesa-
moides, with a decline in phytochemicals observed when the
plants approached their reproductive stages of ﬂowering and
fruiting. In C. triloba, crude extracts showed the presence of phe-
nolics, phlobatannins, saponins, steroids, alkaloids as well as ter-
penoids, with the absence of ﬂavonoids, tannins, cardiac glyco-
sides and cynogenic glycosides (Akula and Odhav, 2008;Mohanlall
et al., 2011;Mudzwiri, 2007). Currently, few studies have in-
vestigated the phytochemical content of these plant species. Thus,
more studies need to be conducted in order to critically evaluate
and document secondary metabolites synthesized by these plants
in order to fully explore their beneﬁts in pharmacology.
Extraction of C. sesamoides and C. triloba plants has resulted in
the isolation and identiﬁcation of a number of compounds with
signiﬁcant importance in nutrition and pharmacology. Sesamin
and sesamolin compounds have been extracted from the seed of C.
sesamoides (Bedigian, 2003). The aforementioned compounds are
generally isolated from the Sesamum genus and have been studied
for their pharmacological properties (Jeng and Hou, 2005). In
glandular trichomes of C. triloba,Ohkawa et al. (2012) isolated nine
malonylated glycerolipids (Fig. 2). The authors found that mal-
onylated glycerolipids represented approximately 90% of the
glandular trichome cell content in C. triloba, with 1-O-acetyl-2-O-
[(R)-3-acetyloxyicosa-noyl]-3-O-malonylglycerol being the most
abundant constituent (41%) followed by 1-O-acetyl-2-O-[(R)-(3-
Documented studies on the nutrient content of Ceratotheca sesamoides and Ceratotheca triloba.
Scientiﬁc name Plant part
Ceratotheca sesamoides Endl. Leaves The two leaf cultivars of Ceratotheca sesamoides were reported to contain high levels of
protein (29.35–29.85%), calcium (2.60–2.62%) and phosphorus (0.25–0.27%). While the seeds
contained reasonable amount of oil levels (17.25–21.00%), crude protein (21.32–22.15%),
crude ﬁbre (25.75–29.75%), calcium (2.65–3.15%) and phosphorus (0.53–0.54%).
Ceratotheca sesamoides Endl. Leaves At three different growth stages (6, 8 and 10 weeks); the moisture and carbohydrate content
were highest on the 6th week 9.30% and 59.90%, respectively. While the highest protein
(12.85%), ash (8.55%) and crude ﬁbre content (8.60%) were recorded on the 8th growth week.
Fasola and Ogunsola
Ceratotheca sesamoides Endl. Leaves The nutritional content of Ceratotheca sesamoides was recorded as dry matter (%) 93.39,
protein 15.11, lipid 0.09, total sugar 75.43, reductors sugar 8.20 per 100 g
D.M. and energy
Yadang et al. (2009)
Ceratotheca sesamoides Endl. Leaves Leaf extracts of Ceratotheca sesamoides contained: water 81 g, energy 226 kJ (54 kcal), protein
4.2 g, fat 0.5 g, carbohydrate 11.0 g, calcium 300 mg, phosphorus 86 mg, iron 3.2 mg, ascorbic
acid 28 mg per 100 g. The seeds were reported to contain: water 7.0 g, energy 2303 kJ
(550 kcal), protein 14.2 g, fat 46.5 g, carbohydrate 27.5 g, calcium 887 mg, iron 38 mg, thia-
min 0.75 mg, riboﬂavin 0.3 mg and niacin 4.4 mg.
Grubben and Denton
Ceratotheca sesamoides Endl. Leaves The leaf extracts of the species contained: dry matter 84.6%, protein 25.0, fat 4.3, ﬁbre 51.1,
carbohydrates 6.1, ash 13.6 g/100 g dry matter and energy 673 (kJ)/161 (kcal). Inorganic
constituents: iron 146.2, zinc 5.0, calcium 1207, phosphorus 377, potassium 2125, copper 2.6,
manganese 20.7, magnesium 592 mg/100 dry matter.
Mitchikpe et al. (2008)
Ceratotheca triloba (Bernh.)
Leaves Ceratotheca triloba extracts contain: energy 259 kJ (62 kcal), moisture 85 g, protein 2 g, fat
2.1 g, ﬁbre 2.07 g, ash 2.27 g, carbohydrates 8.28 g. Mineral content: calcium 705, phosphorus
223, sodium 115, manganese 8, copper 3, zinc 3, magnesium 428, iron 19 mg/100 g dry
Odhav et al. (2007)
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221 213
Documented studies on the phytochemical content of Ceratotheca sesamoides and Ceratotheca triloba.
Scientiﬁc name Plant part
Extracting solvent (s) Phytochemical bioassay (s) Finding(s) Reference
Leaves ND Flavonoids Leaf extracts showed relatively favourable ﬂavonoid content after
8 weeks (1400 mg/100 g) of growth in comparison to 6 and 10 weeks.
Fasola and Ogunsola
Leaves ND Saponins Ceratotheca sesamoides leaves accumulated more saponins as growth
weeks (6–10) progressed.
Fasola and Ogunsola
Leaves ND Alkaloids Alkaloid content ranged from 825 to 855 mg/100 g with the highest
concentration recorded after 10 weeks of growth.
Fasola and Ogunsola
Leaves ND Tannins There was a decrease in tannin content (130–125 mg/100 g) in leaf
extracts during the 10th week of growth.
Fasola and Ogunsola
Leaves ND Phenols Leaf extracts contained the highest phenol content (48 mg/100 g) after
the 8th week of growth.
Fasola and Ogunsola
Leaves Methanol Phenolics Approximately 186 mg GAE 100 g
of phenolic content was measured
in leaf extracts of Ceratotheca sesamoides.
Konan et al. (2014)
Leaves Water Alkaloids, saponins, ﬂavonoids, phenolics, tannins,
cardiac glycosides, steroids, cardenolides and
Leaf extract were reported to contain alkaloids, saponins, ﬂavonoids
and phenolics with no traces of tannins, cardiac glycosides, steroids,
cardenolides and dienolides.
Toyin et al. (2012)
Leaves Methanol, water Phenolic Methanol extracts showed high phenolic content (35.2 mg/g) when
compared to aqueous extracts (14.6 mg/g).
Akula and Odhav
Roots Water Phlobatannins, saponins, steroids, terpenoids, ﬂa-
vonoids, tannins and cardiac glycosides
Major phytochemical compounds including phlobatannins, saponins,
steroids and terpenoids were detected from crude extracts of Cer-
atotheca triloba with no detection of ﬂavonoids, tannins and cardiac
Mohanlall et al.
Leaves 80% Methanol, 10% acetic acid in
ethanol, water, hydrochloric acid
Saponins, alkaloids, oxalic acid and cyanogenic
Leaf extracts contained saponins (0.58 mg/ml), alkaloids (0.6 g/5 g) and
oxalic acid (0.1275 % w/w) with no detection of cyanogenic glycosides.
ND –Not deﬁned.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221214
acetyloxyoctadecanoyl)-3-O-malonylglycerol (21%). Furthermore,
the glandular trichomes were reported to contain minor com-
pounds with iso- and anteiso-type structures in the 3-acetyloxy-
fatty acyl groups in the fatty acyl moiety (Ohkawa et al., 2012).
Furthermore, C. triloba root extracts were isolated for three new
anthraquinones and steroid androgen, androst-5-ene-3, 17, 19-triol
(Mohanlall et al., 2011). These included 9,10 anthracenedione,
rahydro-1H,4aH-phenanthrene-4,9-dione, 1,2 benzenedicarboxylic
acid, mono(2-ethylhexyl)ester, octadecanoic acid and androst-5-
ene-3,17,19-triol (Fig. 3). Amongst the compounds, authors re-
ported on the antioxidant and antibacterial activities displayed by
9,10 anthracenedione and 1-hydroxy-4-methylanthraquinone with
poor anti-inﬂammatory activity (Table 4). The compounds were
further analysed to be potent inhibitors of the human
topoisomerase II enzyme. The structures of anthraquinones were
found to have common characteristics with that of mitoxanthrone
used in the treatment of related ailments such as prostate cancer,
acute myelogenous leukaemia (AML) and breast cancer. In addi-
tion, anthraquinones (natural and synthetic) have been recognized
by the pharmaceutical industries for their crucial role in anti-
bacterial, antitrypanosomal and antineoplastic activities (Heyman
et al., 2009;Tarus et al., 2002).
7. Pharmacological properties of Ceratotheca species
The increasing resistance of micro-organisms to available an-
tibiotics is a major concern to researchers and clinicians around
the world. Existing drugs are becoming ineffective against viruses,
bacteria, fungi and protozoa which is a problem in the ﬁght against
microbial related ailments. In addition to the search for new drugs
from medicinal plants, leafy vegetables might be a complementary
option because of their health beneﬁts both nutritionally and
medicinally. Furthermore, in vitro and epidemiologic studies have
shown the beneﬁts of foods rich in phenolic compounds in redu-
cing the risk of such health problems due to their antioxidant,
anti-mutagenic, anti-inﬂammatory and antibacterial properties
(Gülçin, 2012;Surh, 2002). The antioxidant, antimicrobial, anti-
diarrhoeal, anti-inﬂammatory, anti-diabetic, antiplasmodial, anti-
viral and antivenom activities of C. sesamoides and C. triloba are
presented in Table 4. Different plant parts and extracting solvents
have been assessed using in vitro,ex vitro and in vivo screening for
7.1. Antioxidant activity
Antioxidant activity was found to be moderate in methanol leaf
extracts of C. sesamoides when evaluated against DPPH (2, 2-di-
phenyl-2-picrylhydrazyl hydrate) free radicals (Konan et al., 2014).
Authors found that there was high correlation coefﬁcients (0.93)
between extract concentration and free radical scavenging activity.
Leaf extracts of C. triloba showed reasonably good antioxidant
activity when compared to the positive (rutin, 100% activity)
control (Odhav et al., 2007). Similar antioxidant activity ( 480%
DPPH) was demonstrated by Akula and Odhav (2008) in methanol
Fig.2. Structures of 1-O-acetyl-2-O-(3-acetyloxy-fatty acyl)-3-O-malonylglycerols
(1–9) isolated from glandular trichomes of Ceratotheca triloba. The methyl esters of
1–9are referred to as 1A–9A, whereas methyl esters of 3-hydroxy-fatty acids ob-
tained by hydrolysis of 1–9are designated as 1a–9a.
9,10 anthracenedione 1-hydroxy-4-methylanthraquinone
1,2 benzenedicarboxylic acid,
Fig. 3. Chemical structures of anthraquinone derivatives isolated from Ceratotheca triloba.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221 215
Documented studies on the pharmacological properties of Ceratotheca sesamoides and Ceratotheca triloba.
Scientiﬁc name Plant part
Extracting solvent (s) Test system and positive control
Bioactivity Report on the activity Reference
Leaves Methanol Ascorbic acid, Gallic acid (ND) Antioxidant –DPPH
The extracts were effective (57.0%) against DPPH radical sca-
vengers with an IC
of 7.5 μgml
Konan et al.
Leaves Methanol, Water Rutin (1 mM) Antioxidant –DPPH
Methanol extracts recorded high (84.9%) activity while the water
extracts showed low (36.7%) activity against DPPH radical
Akula and Odhav
Roots Methanol Quercetin-3-rutinoside (1 mM) Antioxidant –DPPH
Approximately750% scavenging activity was observed from
puriﬁed root extracts of Ceratotheca triloba, with
CTRE 02 extract
recorded to have the highest activity (IC
Leaves Methanol Rutin (ND) Antioxidant –DPPH
Leaf extracts demonstrated high (84%) antioxidant activity. Odhav et al.
Leaves, roots Methanol, water In vitro –Gentamycin (100 μg/ml) Antibacterial agar disk
Leaf extract exhibited inhibitory activity against Bacillus cereus,
Enterobacter aerogenes and Micrococcus luteus. While the root
extract showed inhibitory activity against Bacillus cereus and
Odhav (2013)Ampicillin (100 μg/ml)
Leaves, roots Acetone, water In vitro –Gentamycin (100 μg/ml) Antibacterial agar disk
The six isolated compounds showed good activity against the
Gram positive bacteria Micrococcus luteus, Bacillus cereus and
Staphylococcus aureus. Whereas the Gram negative bacteria ex-
hibited moderate activity against Escherichia coli and Salmonella
Odhav (2013)Ampicillin (100 μg/ml)
Leaves, roots Methanol, water In vitro –Amphotericin B (5 mg/ml) Antifungal activity There was minimum antifungal activity observed for the two
strains of Aspergillus ﬂavus and Fusarium verticilloides fungi.
Plant Saline In vitro –Human epidermoid carci-
noma HEP-2 cell line (ND)
Antiviral activity Extracts exhibited a dose dependent activity against measles
virus (HEP-2). Virusþcell þextract showed antiviral activity at 10
and 15 mg/ml extract concentration. While the cell treated with
an extract and virus exerted a mild antiviral activity at 15 mg/ml
Obi et al. (2006)
Leaves Water Ex vitro –Loperamide Antidiarrhoeal activity Aqueous leaf extract of Ceratotheca sesamoides showed a dose
dependent activity. Extracts at 25 mg/kg body weight sig-
niﬁcantly (194.50 min) prolonged the onset time of diarrhoea in
rats. While there were no signs of diarrhoea in rats treated with
50 and 100 mg/kg body weight of the extracts.
Toyin et al. (2012)
Hydrochloride (1 ml)
Leaves Methanol In vitro –Nordihydroguaiaretic acid
Plant extracts exhibited minimum inhibitory activity (IC
56.1 μg/ml) towards the 5-LOX enzyme.
Akula and Odhav
Rutin (7.3 μg/ml)
Leaves Methanol In vitro –Nordihydroguaretic acid
Ceratotheca triloba leaves and roots showed poor (IC
ml) anti-inﬂammatory activity against the 5-LOX enzyme.
Leaves Water In vitro –Acarbose (1 mg/ml) Anti-diabetic activity Aqueous extract had a dose dependent response in the inhibition
of α-amylase (1 mg/ml: 43.23%; 3 mg/ml: 99.31%; 5 mg/ml:
Odhav et al.
Whole plant Petroleum ether, acetone,
In vitro –Chloroquine (400 ng/ml) Antiplasmodial activity Plant extracts had an IC
ranging from 20 to 450 mg/ml in both
fresh and stored material against Plasmodium falciparum (FcM29-
Benoit-Vical et al.
Whole plant Petroleum ether, acetone,
In vitro –Chloroquine (60 ng/ml) Antiplasmodial activity In the Plasmodium falciparum (FcB1-Colombia strain), extracts
had an IC
ranging from 4 to 450 mg/ml. With the best activity
observed from stored material extracted with ethanol.
Benoit-Vical et al.
Leaves Water, dichloromethane/
Ex vitro –N,N-diethyl-meta-
Antiplasmodial activity Dichloromethane/Methanol extracts were reported to be 50%
effective against Anopheles arabiensis in rats. While water ex-
tracts showed 30% Anopheles arabiensis inhibition.
Maharaj et al.
Twigs Water, dichloromethane/
Ex vitro –N,N-diethyl-meta-
Antiplasmodial activity Dichloromethane/methanol and water extracts had a repellent
effect of 40% and 31%, respectively against Anopheles arabiensis in
Maharaj et al.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221216
extracts. However, leaf aqueous extracts in the aforementioned
study recorded low activity against DPPH radical scavengers. In
Table 4, puriﬁed root extracts (9,10 anthracenedione and 1 hydro-
xy-4-methyl anthraquinone) of C. triloba showed approximately
50% DPPH antioxidant activity (Mohanlall and Odhav, 2013). In
order to understand the moderate antioxidant activity demon-
strated by puriﬁed root extracts of C. triloba, crude root extracts
need to be investigated. From the documented studies, C. sesa-
moides and C. triloba showed moderate–good activity against the
DPPH free radicals, however more studies need to be conducted in
order to determine the β-carotene activity of both species. Studies
have suggested that choosing at least two antioxidant bioassays
such as DPPH and β-carotene is critical due to their complex
processes (Moon and Shibamoto, 2009).
7.2. Antimicrobial and antiviral activity
In the in vitro agar disk diffusion bioassay, leaves of C. triloba
exhibited inhibitory activity against Gram-positive bacteria (Ba-
cillus cereus and Micrococcus luteus) and a Gram-negative bacter-
ium (Enterobacter aerogenes)(Mohanlall and Odhav, 2013). On the
other hand, root extracts were effective against B. cereus and
M. luteus. The authors further reported on the inhibitory activity of
9,10 anthracenedione against Escherichia coli and Salmonella ty-
phimurium as well as 1-hydroxy-4-methylanthraquinone against
Staphylococcus aureus and M. luteus.McGaw et al. (2000) found
that C. triloba exhibited poor antibacterial activity when tested
against Gram-positive bacteria (B. subtilis and S. aureus) and Gram-
negative bacteria (E. coli and Klebsiella pneumoniae). In order to
avoid such conﬂicting results, more studies need to be conducted
to investigate the antibacterial properties of the species. Further-
more, the lack of information on the antimicrobial properties of C.
sesamoides is of concern since the plant is traditionally used for
microbial-related ailments such as conjunctivitis and leprosy.
Antiviral activity of C. sesamoides plant extracts was evaluated
in using in vitro assays (Obi et al., 2006). The extracts were effec-
tive against the measles virus on human epidermoid carcinoma
HEP-2 cell line when tested at high concentrations (10 and 15 mg/
ml). Good activity was observed when C. sesamoides extracts
(highest concentration) were treated with the virus and cells. The
authors suggested on the use of C. sesamoides against the measles
virus since in Nigeria it is utilized in the management of con-
junctivitis, a viral infection. Findings from this study should be a
stepping stone in the search for antiviral drugs as well as other
viral related diseases that are a major concern worldwide.
7.3. Antidiarrhoeal activity
Castor oil induces problems such as water and electrolyte
permeability, resulting in a change in the intestinal mucosal
membrane that lead to ﬂuids and watery luminal content that ﬂow
rapidly through the small and large intestines causing diarrhoea.
When the antidiarrhoeal properties of C. sesamoides were eval-
uated ex vitro using castor oil, aqueous leaf extracts inhibited the
onset of diarrhoea at concentrations of 50 and 100 mg/kg plant
extract body weight in rats (Toyin et al., 2012). At these con-
centrations, the typical diarrhoeal symptoms were not noticeable
in the experimental rats. Most importantly, the extracts showed
almost similar potency (194.50 min) as that of the positive control
(Loperamide hydrochloride, 233 min) when tested at the lowest
concentration (25 mg/kg body weight). C. sesamoides showed po-
tential in the inhibition (55.56% defecation inhibition) of diarrhoea
at the lowest extract concentration, similar to the positive control
(Toyin et al., 2012). With a 100% defecation inhibition observed in
plant extract at 50 and 100 mg/kg body weight in rats. The authors
suggested that diarrhoea may have been prevented by the activity
Fruits Water Ex vitro –N,N-diethyl-meta-
Antiplasmodial activity Water extracts recorded a 26% Anopheles arabiensis inhibition in
Maharaj et al.
Herb Water, ethanol In vitro –Aristolochic acid (ND) Hyaluronidase activity Water extracts recorded higher hyaluronidase inhibition (14%)
compared to the ethanol extracts (1%) for the Bitis arietans ve-
nom. However, better hyaluronidase inhibition was observed
from water and ethanol extracts against Naja nigricollis venom
with 50% and 27%, respectively.
Molander et al.
Herb Water, ethanol In vitro –EDTA (ND) Phospholipase A
activity Aqueous extracts were reported to have 27% phospholipase A
inhibition while ethanol extracts had 10% when tested against
Bitis arietans venom.
Molander et al.
Herb Water, ethanol In vitro –4-(2-Aminoethyl)benzene-
sulfonyl ﬂuoride, EDTA (ND)
Proteolytic activity Both solvent extracts showed very low ( r4) proteolytic inhibi-
tion against Bitis arietans venom.
Molander et al.
DPPH (2, 2-diphenyl-2-picrylhydrazyl hydrate) bioassay, ND –not deﬁned.
(i) 9,10 anthracenedione, (ii) 1-hydroxy-4-methylanthraquinone, (iii) 5,8-dimethoxy-2,3,10,10a-tetrahydro-1H,4aH-phenanthrene-4,9-dione, androst-5-ene-3,17,19-triol, (iv) 1,2 benzenedicarboxylic acid, (v) mono (2-ethyl-
hexyl) ester (vi) octadecanoic; CTRE 01: 9,10 anthracenedioneþ1-hydroxy-4-methylanthraquinone; CTREh02 and CTREh03; HEP-2 cell line-Human epidermoid carcinoma HEP-2 cell line.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221 217
of C. sesamoides on prostanglandin synthesis, nitric oxide and
platelet activating factor production, since prostaglandin inhibitors
and nitric oxide syntheses have been attributed to the delay of
diarrhoea induced by castor oil in rats.
7.4. Anti-inﬂammatory activity
Inﬂammation results from excess generation of reactive oxygen
species (ROS) stimulated by the production of cytokines and en-
zyme activation such as lipoxygenases (LOXs) from inﬂammatory
cells. The LOX enzyme is responsible for several inﬂammation
related diseases (Dobrian et al., 2011). Lipoxygenases is involved in
the biosynthesis of leukotrienes and prostagladins which if in-
hibited, results in the prevention of several diseases triggered by
oxidative stress (Rådmark and Samuelsson, 2007). Regulation of
ROS production is an important step in the down-regulation of the
immune response thus preventing chronic inﬂammation (Seifried
et al., 2007).
Table 4 represents the in vitro inhibition of 5-lipoxygenase (5-
LOX) enzyme activity in C. triloba leaf and root extracts. Methanol
leaf extracts exhibited minimal inhibitory effects against the 5-LOX
enzyme with an IC
of 56.1 mg/ml (Akula and Odhav, 2008). Simi-
larly, Mohanlall and Odhav (2013) reported on the poor anti-in-
ﬂammatory activity of leaves and root extracts (IC
of 300 mg/ml)
when tested against the 5-LOX enzyme. The poor activity of C. tri-
loba is somehow contradictory to its traditional use in pain-related
ailments such as stomach cramps and painful menstruation. Such
insigniﬁcant/poor anti-inﬂammatory activity of plant extracts
against the 5-LOX enzyme might be due to several factors including
the tested enzyme. According to Jäger et al. (1996),activecom-
pounds of the extracts might be more effective when tested at other
sites in the complex process of inﬂammation. Therefore, more
studies examining the activity of C. triloba against LOX enzymes and
other enzymes such as cyclooxygenase-1 and 2 are required.
7.5. Anti-diabetic activity
Traditionally, there has been no record of C. triloba in the
treatment of diabetes. Thus, the inhibitory activity of leaf extracts
against the α-amylase enzyme at the lowest (1 mg/ml) and highest
(5 mg/ml) concentrations recorded to be 43.3% and 99.7%, re-
spectively is noteworthy (Odhav et al., 2010). The inhibitory ac-
tivity of C. triloba extracts at 3 and 5 mg/ml was equivalent to that
of the positive control with a 99.2% inhibitory activity (acarbose at
1 mg/ml). Therefore, ﬁndings from Odhav et al. (2010) might
contribute to the search for plants with anti-diabetic properties.
Furthermore, these ﬁndings are of interest since the plant species
will not only provide nutritional beneﬁts but also play a vital role
in inhibiting the α-amylase enzyme, thereby reducing blood glu-
7.6. Antiplasmodial activity
The in vitro and ex vitro antiplasmodial properties of C. sesa-
moides and C. triloba are presented in Table 4. Plant extracts of C.
sesamoides were more effective after storage for the Plasmodium
falciparum (FcB1-Colombia) strain (Benoit-Vical et al., 2008).
Amongst the solvents used, ethanol extracts gave good activity
–4mg/ml) compared to petroleum ether (IC
and acetone (IC
–33 mg/ml). Overall, plant extracts (fresh or
stored) were more effective against the FcB1-Colombia strain in
comparison to the FcM29-Cameroon strain. In C. triloba plants,
dichloromethane/methanol leaf extracts yielded the highest (50%)
efﬁcacy against Anopheles arabiensis in rats after 2 min during ex
vitro studies (Maharaj et al., 2010). While the antiplasmodial in-
hibitory activity of other plant parts such as twigs and leaves
ranged from 26% to 40% when extracted with, either di-
chloromethane/methanol or water. This study has revealed the
difference in the plants ability to inhibit various malarial strains,
thus more strains need to be investigated. These results also justify
the traditional use of C. sesamoides in the treatment of malaria.
7.7. Hyaluronidase, phospholipase and proteolytic activity
Since C. sesamoides is traditionally used against snakebites, it
was evaluated for plant compounds effective against the necrosis-
inducing enzymes of snake venom (Molander et al., 2014). Plant
extracts (5 mg/ml) showedr50% activity against the tested en-
zymes of Bitis arietans (5 mg/ml) and Naja nigricollis (5 mg/ml). In
most cases, water extracts were found to be more potent in in-
hibiting hyaluronidase, phospholipase and proteolytic enzymes
from B. arietans and N. nigricollis venom compared to ethanol
extracts. From the N. nigricollis venom, water extracts demon-
strated the highest (50%) hyaluronidase enzyme inhibition in
comparison to the ethanol extracts. The ability of C. sesamoides to
inhibit some enzymes found in snake venom might be due to the
polyphenols such as tannins (Molander et al., 2014). The ability of
aqueous extracts to inhibit 50% of hyaluronidase enzymes from N.
nigricollis venom is of interest, thus more extract concentrations
should be evaluated.
7.8. Other documented properties
Table 5 represents other biological activities of C. sesamoides. In
vivo insecticidal activity was investigated with C. sesamoides seeds
(Uddin II and Abdulazeez, 2013). Seed extracts (powdered and
aqueous treatments) exhibited high mortality rate (3.00) against
Callosobruchus maculatus at the lowest tested concentration (1.5%)
after 72 h in infested cowpea seed. In comparison to the negative
control (water), seed extracts prevented more oviposition of C.
maculatus even at the lowest extract concentration. C. sesamoides
has been reported to be pest tolerant (Grubben and Denton, 2004)
and has been found to be visited by 149 insects with minimal
damage observed on the plant (Uddin and Adesiyun, 2011). The
potential organic pesticide of the plant against cowpea seeds
needs to be further evaluated especially the secondary metabolic
compounds synthesized by the plant to cause it to be pest tolerant.
Such ﬁndings may give an understanding on the insecticidal ac-
tivity of C. sesamoides since more studies are now focused on
‘green farming’with organic fertilizers becoming more popular
(Aguilera et al., 2013). Furthermore, studies evaluating volatile
compounds in C. sesamoides are pertinent since the plant has an
acrid smell which might be the reason for its insecticidal activity.
The aim of the phytochemistry (Table 3) and pharmacological
section (Tables 4 and 5) was to understand the potential of the
plant species in the treatment of several ailments as well as to
show the limited literature information available. In the phy-
tochemistry section, few studies have focused on the isolation and
identiﬁcation of compounds (Figs. 2 and 3)inCeratotheca plant
species (Bedigian, 2003;Ohkawa et al., 2012;Mohanlall et al.,
2011). From the available studies, the plant species yielded a wide
range of compounds that can be useful in the nutritional and
pharmacological sectors. These compounds are of value especially
the anthraquinones isolated from C. triloba which are recognized
for their similarities with mitoxanthrone compounds used for
cancer related diseases. Nevertheless, studies directed towards the
isolation and identiﬁcation of compounds are of necessity. In ad-
dition, investigation of these compounds is crucial if they are to be
utilized in pharmacological industries.
In the current review, Ceratotheca species indicated potential
antimicrobial, antiviral, antidiarrhoeal, anti-diabetic and anti-
plasmodial properties. Although the species have been
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221218
documented in the treatment of a wide range of microbial ail-
ments in traditional medicine, their antimicrobial properties for
crude plant extracts were very poor (McGaw et al., 2000) com-
pared to isolated compounds (Mohanlall and Odhav, 2013). Diar-
rhoea is caused by a wide range of entero-pathogenic micro-
organisms such as Shigella ﬂexneri,S. aureus,E. coli,Salmonella
typhi and Candida albicans. The antidiarrhoeal activity demon-
strated by C. sesamoides was noteworthy (Toyin et al., 2012). The
results from the study serves to conﬁrm the claims made in tra-
ditional medicine about the plant being used in the treatment of
diarrhoea. Furthermore, C. sesamoides plant extracts were reported
to inhibit the measles virus on the human epidermoid carcinoma
HEP-2 cell line (Obi et al., 2006). The ability of Ceratotheca species
to inhibit microbial and viral infections is of interest due to the
increasing resistance of microorganisms to available antibiotics
worldwide. Even though C. triloba showed minimal inhibitory ef-
fects against the 5-LOX enzyme, the plant extracts was effective
against the α-amylase enzyme. However, further isolation and
identiﬁcation of the active hypoglycaemia and hypolipidemic
principles in the plants are needed in order to understand their
mechanisms of action. C. sesamoides plant extracts were also re-
ported to have antiplasmodial and insecticidal activities. The low
potency of fresh plant material against Plasmodium strains is of
interest since most plants in traditional medicine are dried,
ground and stored before usage.
Besides the inadequate literature on the biological activities of
Ceratotheca species, few studies have focused on investigating the
effect of plant parts (crude or isolated compounds), solvent ex-
tracts and extract dose/concentration. From the available studies,
leaves were the most commonly used plant part compared to
roots, seeds and the whole plant from both species. The potency of
leaf extracts in the evaluated biological activities is important for
conservational purposes of Ceratotheca species from wild popula-
tions. Generally, water extracts are known to exhibit lower activity
when compared to non-polar extracts in various pharmacological
studies. Nevertheless, aqueous extracts of Ceratotheca species
showed good antidiarrhoeal, anti-diabetic, hyaluronidase enzyme
and insecticidal activities. Ceratotheca species showed diverse
bioassay activities including in vitro,ex vitro and in vivo. From
these studies, it is evident there is still a dire need for more re-
search especially for both species since they might have similar
potency against various diseases. Once pharmacological potential
of plants has been established in in vitro and in vivo bioassays,
isolation and investigation of the compounds will be necessary.
8. Safety and toxicity of Ceratotheca species
Although Ceratotheca species play a major role as food sup-
plement in most diets (Table 2), the species are also widely used
for their medicinal properties in most African countries (Table 4).
Therefore, it is crucial to determine their safety and toxicity
especially since the plants are consumed in large quantities (Ta-
ble 5). In a test to determine toxicity of C. sesamoides, the extracts
showed minimal toxicity against the vero cell line with acetone
extracts having no cytotoxic effect even at high tested concentra-
tions (Benoit-Vical et al., 2008). Puriﬁed extracts of C. triloba ex-
hibited cytotoxic activity (41.4–73.2%) against HepG
cells at high
concentrations (Mohanlall, 2010;Mohanlall and Odhav, 2013).
Nevertheless, low puriﬁed extract concentration (1.56 mg/ 25 ml)
showed slight cytotoxicity against HepG
cells. At 1000 mg/ml,
organic extracts of C. triloba demonstrated low levels of cytotoxi-
city withr5% cell death in HepG
cells while aqueous extracts
were non-cytotoxic (Mudzwiri, 2007). Toxic analysis of C. triloba
showed that the plant extracts did not contain cyanogenic glyco-
sides (Mudzwiri, 2007). However, aqueous extracts were
Other documented pharmacological activities of Ceratotheca sesamoides and Ceratotheca triloba.
Species Plant part
Extracting solvent (s) Test system and positive control
Bioactivity Report on the activity References
Seeds Water In vivo –Pirimiphos-methyl
Insecticidal activity In both extracts, Callosobruchus maculatus mortality was highest at 1.5%
concentration after 72 h of cowpea infestation. Aqueous extracts pre-
vented oviposition and F
progeny emergence of Callosobruchus macu-
latus at lower concentration compared to powder extracts.
Uddin and Abdulazeez
Plant Petroleum ether, acet-
In vitro –Etoposide
Plant extracts showed slight toxicity at high concentration. Benoit-Vical et al.
Leaves Water, 80% methanol In vitro –ND Cytotoxicity
Methanol extracts had minimal cytotoxicity with the aqueous extracts
exhibiting no cytotoxic effect.
Leaves, roots Methanol, water In vitro –ND Cytotoxicity
Puriﬁed extracts of Ceratotheca triloba had 41.4–73.2% HepG
cell death at
12.5 mg/25 ml while low concentrations (1.56 mg/ 25 ml) exhibited 1–14.4%
ND –Not deﬁned.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209–221 219
considered toxic when consumed at levels above 100 and 1000 mg/
ml (Mudzwiri, 2007). C. triloba extracts showed no mutagenic
activity when evaluated in the Ames test (Mohanlall and Odhav,
2013;Mudzwiri, 2007). The species (crude and puriﬁed extracts)
were also reported to have inhibitory activity against the topoi-
somerase II enzyme. Overall, C. sesamoides and C. triloba were
reported to be potentially cytotoxic and toxic when tested at high
concentrations. Thus, more stringent studies need to be conducted
in order to understand the toxicity levels of these leafy vegetables
since they are consumed in high quantities.
9. Propagation strategies of Ceratotheca species
Leafy vegetables are known to form a substantial part of diets
and are often considered the cheapest and most readily available
source of food. Nevertheless, their conservation has become a
concern in international agricultural research sectors. Thus, stra-
tegies combining the standard ex situ and in situ conservation for
indigenous vegetables need to be implemented in order to pro-
mote and improve their utilization. These vegetables are known to
be resistant, adaptable and tolerant to different harsh climatic
conditions compared to exotic species (Raghuvanshi and Singh,
2001). To date, few studies have documented propagation strate-
gies for C. sesamoides and C. triloba.
C. sesamoides and C. triloba are relatively easy to cultivate.
Plants can be propagated through seeds which are sown at the
onset of the rainy season during early summer. Plants grow best in
rich well-drained sandy soils with full sun exposure or semi-shade
(Duncan, 2011;Grubben and Denton, 2004). Seedlings of C. triloba
are often slow to grow initially. C. sesamoides seeds do not show
dormancy and can be intercropped with okra, eggplant, cowpea,
amaranth, sorghum, sweet potato and sesame (Grubben and
Denton, 2004). The species are drought, disease and pest tolerant.
Frequent pruning of shoots permits sustained vegetative growth
and ﬂowering, prolonging the growth cycle.
In an effort to improve the mutagenesis of C. sesamoides,Nura
et al. (2014) found that 0.1 mM colchicine concentration was ef-
fective in the genetic improvement of different growth para-
meters. According to the authors, improved growth parameters
were as a result of the plants ability to respond well to the mu-
tagens (colchicine) in inducing favourable mutants. Similarly,
0.1 mM colchicine concentration was reported to increase the
overall yields of cultivated C. sesamoides plants (Nura et al., 2012).
Fasakin and Oloﬁntoye (2005) established that for different culti-
vars, row spacing as well as seeding rate had a positive effect on
plant population density in C. sesamoides while the overall yields
were improved by different cultivar and seeding rates.
To date, few studies have evaluated the nutritional and med-
icinal role of the under-utilized leafy vegetables, C. sesamoides and
C. triloba. This might be attributed to several reasons such as the
preference of other leafy vegetables, their availability and the
plants being considered as “poor man’s food”. Nonetheless, the
current review has revealed the nutritional potential (relatively
high energy levels in kilojoules, carbohydrates, protein, fat content
and inorganic constituents) of both species. The current review
shows that utilization of these leafy vegetables could play a crucial
role in dietary diversiﬁcation, improved nutrition and human
health. However, more work is still needed to further validate the
vegetable’s nutritional role if they are to supplement the starch
based diets of African communities. Research also needs to high-
light the variations in the chemical compositions of Ceratotheca
leafy vegetables especially the change in nutritional content
caused by environmental/growth conditions, harvest stages, their
preparation and preservation.
Traditional vegetables like C. sesamoides and C. triloba are fre-
quently categorized as functional foods due to their health beneﬁts
extending beyond basic nutrition (Van Wyk and Gericke, 2000). C.
sesamoides and C. triloba showed potentially good antibacterial,
anti-diarrhoeal, anti-diabetic, antiplasmodial and antiviral proper-
ties against various tested micro-organisms. These properties might
be due to the different secondary metabolites (ﬂavonoids, alkaloids,
saponins, tannins, phenolics and steroids) synthesized by these
leafy vegetables. Most importantly, C. sesamoides and C. triloba were
reported to yield various compounds including sesamin, sesamolin,
anthraquinones similar to mitoxanthrone (for cancer treatment)
and malonylated glycerolipids. From the available information on
the pharmacological role of Ceratotheca species, there is still a dire
need for studies to evaluate the plant’sbiologicalactivitiesinin
vitro and in vivo bioassays, different plant parts, solvent extracts and
extract dose/concentration. Furthermore, more work is still required
in order to elucidate the plants phytochemistry, their relation with
the plants traditional use and their signiﬁcance in human health
especially with the compounds they produce. Such work can also
assess the quantity and up-regulation of important compounds
such as anthraquinones for their exploitation. It is vital that more
research is conducted on potentially exploitable wild plant species
such as Ceratotheca. This would promote their increased utilization
thereby simultaneously contributing to conserving their genetic
resources. In order to introduce the species as a conventional ve-
getable crop as well as a medicinal plant, renewed efforts aimed at
cultivation practices are pertinent. Safety and toxicity analysis of C.
sesamoides and C. triloba remains a major concern and needs to be
extensively examined especially since the plants are consumed in
Financial support from the National Research Foundation,
Pretoria, Graça Machel (Canon Collins) and the University of
KwaZulu-Natal (Pietermaritzburg), South Africa is appreciated. Dr.
J.J. Nair is thanked for his assistance in the compilation of isolated
chemical structures. Dr. M. Moyo is thanked for his suggestions on
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