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Nutritional and pharmacological potential of the genus Ceratotheca–An underutilized leafy vegetable of Africa

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
article info
Article history:
Received 25 August 2015
Received in revised form
30 November 2015
Accepted 1 December 2015
Available online 4 December 2015
Nutrient analysis
Phytochemical content
Propagation strategies
Traditional leafy vegetables
Ethnopharmacological relevance: Ceratotheca (Pedaliaceae) is an endemic African genus comprising of
ve 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-inammatory activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
7.5. Anti-diabetic activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
7.6. Antiplasmodial activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
7.7. Hyaluronidase, phospholipase and proteolytic activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
Contents lists available at ScienceDirect
journal homepage:
Journal of Ethnopharmacology
0378-8741/&2015 Elsevier Ireland Ltd. All rights reserved.
Corresponding author.
E-mail address: (J. Van Staden).
Journal of Ethnopharmacology 178 (2016) 209221
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
1. Introduction
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 ve 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 ve 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 classied 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-
ammatory 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
classied as functional foods due to their health benets 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 signicance of the African endemic species
C. sesamoides and C. triloba, and their nutritional and pharmaco-
logical properties.
Fig. 1. Ceratotheca morphology: Awhole plant; Bowers; and Cseed capsule.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209221210
2. Botanical distribution and general morphology of Cer-
atotheca species
The ve 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
0.62.0 m. Ceratotheca leaves are opposite with lower leaves larger
and the uppermost leaves narrowly to broadly lanceolate. The
owers 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 sesamewas rst de-
scribed by Prof. Stephan Endlicher (18041849), 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 rst
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 Curtiss
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 owers 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
conservation purposes.
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 insufcient (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 insignicant 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;NDanikou 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 signicant 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
classied as a poor mans 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 mans 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, bres 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) 209221 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, owers and
seeds. C. sesamoides is prepared by grinding leaves into ne
powders and mixing them with groundnut our, 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 specic
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 owers 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 (0.210.45 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 (45 fold) in comparison to its addition
which results in decreased iron absorption (Hurrell et al., 1992).
Table 1
Documented surveys on the consumption of Ceratotheca sesamoides and Ceratotheca triloba in African countries.
Scientic name Distribution Status of distribution Result highlights References
Ceratotheca sesamoides Endl. Benin (SudanoGuinean and Suda-
nian region)
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.
NDanikou et al. (2011)
Ceratotheca triloba (Bernh.)
Zimbabwe Wild In the 32 edible traditional vegetables identied in Zimbabwe, Ceratotheca triloba was found to be
consumed in Beitbridge, Binga, Tanganda Halt and Umguza Districts. It has been classied as a poor
mans foodtherefore not traded in the markets.
Maroyi (2011)
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209221212
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 scientic 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
as food.
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 inamed 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 atulence (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 quantication of C. se-
samoides leaves yielded avonoids, 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 signicant impact on the phytochemical content of C. sesa-
moides, with a decline in phytochemicals observed when the
plants approached their reproductive stages of owering 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 avonoids, 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 benets in pharmacology.
Extraction of C. sesamoides and C. triloba plants has resulted in
the isolation and identication of a number of compounds with
signicant 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-
Table 2
Documented studies on the nutrient content of Ceratotheca sesamoides and Ceratotheca triloba.
Scientic name Plant part
Finding(s) Reference
Ceratotheca sesamoides Endl. Leaves The two leaf cultivars of Ceratotheca sesamoides were reported to contain high levels of
protein (29.3529.85%), calcium (2.602.62%) and phosphorus (0.250.27%). While the seeds
contained reasonable amount of oil levels (17.2521.00%), crude protein (21.3222.15%),
crude bre (25.7529.75%), calcium (2.653.15%) and phosphorus (0.530.54%).
Fasakin (2004)
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 bre 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
(kJ) 1515.75.
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, riboavin 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, bre 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, bre 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) 209221 213
Table 3
Documented studies on the phytochemical content of Ceratotheca sesamoides and Ceratotheca triloba.
Scientic name Plant part
Extracting solvent (s) Phytochemical bioassay (s) Finding(s) Reference
Ceratotheca sesamoides
Leaves ND Flavonoids Leaf extracts showed relatively favourable avonoid content after
8 weeks (1400 mg/100 g) of growth in comparison to 6 and 10 weeks.
Fasola and Ogunsola
Ceratotheca sesamoides
Leaves ND Saponins Ceratotheca sesamoides leaves accumulated more saponins as growth
weeks (610) progressed.
Fasola and Ogunsola
Ceratotheca sesamoides
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
Ceratotheca sesamoides
Leaves ND Tannins There was a decrease in tannin content (130125 mg/100 g) in leaf
extracts during the 10th week of growth.
Fasola and Ogunsola
Ceratotheca sesamoides
Leaves ND Phenols Leaf extracts contained the highest phenol content (48 mg/100 g) after
the 8th week of growth.
Fasola and Ogunsola
Ceratotheca sesamoides
Leaves Methanol Phenolics Approximately 186 mg GAE 100 g
of phenolic content was measured
in leaf extracts of Ceratotheca sesamoides.
Konan et al. (2014)
Ceratotheca sesamoides
Leaves Water Alkaloids, saponins, avonoids, phenolics, tannins,
cardiac glycosides, steroids, cardenolides and
Leaf extract were reported to contain alkaloids, saponins, avonoids
and phenolics with no traces of tannins, cardiac glycosides, steroids,
cardenolides and dienolides.
Toyin et al. (2012)
Ceratotheca triloba
(Bernh.) Hook.f.
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
Ceratotheca triloba
(Bernh.) Hook.f.
Roots Water Phlobatannins, saponins, steroids, terpenoids, a-
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 avonoids, tannins and cardiac
Mohanlall et al.
Ceratotheca triloba
(Bernh.) Hook.f.
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.
Mudzwiri (2007)
ND Not dened.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209221214
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,
1-hydroxy-4-methylanthraquinone, 5,8-dimethoxy-2,3,10,10a-tet-
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-inammatory 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 ght 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 benets both nutritionally and
medicinally. Furthermore, in vitro and epidemiologic studies have
shown the benets of foods rich in phenolic compounds in redu-
cing the risk of such health problems due to their antioxidant,
anti-mutagenic, anti-inammatory and antibacterial properties
(Gülçin, 2012;Surh, 2002). The antioxidant, antimicrobial, anti-
diarrhoeal, anti-inammatory, 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
Ceratotheca species.
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 coefcients (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
(19) isolated from glandular trichomes of Ceratotheca triloba. The methyl esters of
19are referred to as 1A9A, whereas methyl esters of 3-hydroxy-fatty acids ob-
tained by hydrolysis of 19are designated as 1a9a.
9,10 anthracenedione 1-hydroxy-4-methylanthraquinone
1,2 benzenedicarboxylic acid,
octadecanoic acid
Fig. 3. Chemical structures of anthraquinone derivatives isolated from Ceratotheca triloba.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209221 215
Table 4
Documented studies on the pharmacological properties of Ceratotheca sesamoides and Ceratotheca triloba.
Scientic name Plant part
(s) tested
Extracting solvent (s) Test system and positive control
Bioactivity Report on the activity Reference
Ceratotheca sesamoides
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.
Ceratotheca triloba
(Bernh.) Hook.f.
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
Ceratotheca triloba
(Bernh.) Hook.f.
Roots Methanol Quercetin-3-rutinoside (1 mM) Antioxidant DPPH
Approximately750% scavenging activity was observed from
puried root extracts of Ceratotheca triloba, with
CTRE 02 extract
recorded to have the highest activity (IC
1 mg/ml).
Mohanlall and
Odhav (2013)
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves Methanol Rutin (ND) Antioxidant DPPH
Leaf extracts demonstrated high (84%) antioxidant activity. Odhav et al.
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves, roots Methanol, water In vitro Gentamycin (100 μg/ml) Antibacterial agar disk
diffusion activity
Leaf extract exhibited inhibitory activity against Bacillus cereus,
Enterobacter aerogenes and Micrococcus luteus. While the root
extract showed inhibitory activity against Bacillus cereus and
Micrococcus luteus.
Mohanlall and
Odhav (2013)Ampicillin (100 μg/ml)
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves, roots Acetone, water In vitro Gentamycin (100 μg/ml) Antibacterial agar disk
diffusion activity
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
Mohanlall and
Odhav (2013)Ampicillin (100 μg/ml)
Ceratotheca triloba
(Bernh.) Hook.f.
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 avus and Fusarium verticilloides fungi.
Mohanlall and
Odhav (2013)
Ceratotheca sesamoides
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
extract concentration.
Obi et al. (2006)
Ceratotheca sesamoides
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-
nicantly (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)
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves Methanol In vitro Nordihydroguaiaretic acid
(4.1 μg/ml)
Plant extracts exhibited minimum inhibitory activity (IC
56.1 μg/ml) towards the 5-LOX enzyme.
Akula and Odhav
Rutin (7.3 μg/ml)
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves Methanol In vitro Nordihydroguaretic acid
Ceratotheca triloba leaves and roots showed poor (IC
300 μg/
ml) anti-inammatory activity against the 5-LOX enzyme.
Mohanlall and
Odhav (2013)
Roots Water
Ceratotheca triloba
(Bernh.) Hook.f.
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:
99.07%) enzyme.
Odhav et al.
Ceratotheca sesamoides
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-
Cameroon strain).
Benoit-Vical et al.
Ceratotheca sesamoides
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.
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves Water, dichloromethane/
methanol (1:1)
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.
Ceratotheca triloba
(Bernh.) Hook.f.
Twigs Water, dichloromethane/
methanol (1:1)
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) 209221216
extracts. However, leaf aqueous extracts in the aforementioned
study recorded low activity against DPPH radical scavengers. In
Table 4, puried 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 puried root extracts of C. triloba, crude root extracts
need to be investigated. From the documented studies, C. sesa-
moides and C. triloba showed moderategood 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 conicting 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 uids and watery luminal content that ow
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
Ceratotheca triloba
Fruits Water Ex vitro N,N-diethyl-meta-
Antiplasmodial activity Water extracts recorded a 26% Anopheles arabiensis inhibition in
Maharaj et al.
Ceratotheca sesamoides
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.
Ceratotheca sesamoides
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.
Ceratotheca sesamoides
Herb Water, ethanol In vitro 4-(2-Aminoethyl)benzene-
sulfonyl uoride, 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 dened.
(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) 209221 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-inammatory activity
Inammation results from excess generation of reactive oxygen
species (ROS) stimulated by the production of cytokines and en-
zyme activation such as lipoxygenases (LOXs) from inammatory
cells. The LOX enzyme is responsible for several inammation
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 inammation (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-
ammatory 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
insignicant/poor anti-inammatory 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 inammation. 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, ndings from Odhav et al. (2010) might
contribute to the search for plants with anti-diabetic properties.
Furthermore, these ndings are of interest since the plant species
will not only provide nutritional benets but also play a vital role
in inhibiting the α-amylase enzyme, thereby reducing blood glu-
cose levels.
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
15 mg/ml)
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%)
efcacy 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 ndings may give an understanding on the insecticidal ac-
tivity of C. sesamoides since more studies are now focused on
green farmingwith 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
identication 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 identication 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) 209221218
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 exneri,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 conrm 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
identication 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). Puried extracts of C. triloba ex-
hibited cytotoxic activity (41.473.2%) against HepG
cells at high
concentrations (Mohanlall, 2010;Mohanlall and Odhav, 2013).
Nevertheless, low puried 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
Table 5
Other documented pharmacological activities of Ceratotheca sesamoides and Ceratotheca triloba.
Species Plant part
(s) tested
Extracting solvent (s) Test system and positive control
Bioactivity Report on the activity References
Ceratotheca sesamoides
Seeds Water In vivo Pirimiphos-methyl
(0.5% w/w)
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
Ceratotheca sesamoides
Plant Petroleum ether, acet-
one, ethanol
In vitro Etoposide
(2 10
Plant extracts showed slight toxicity at high concentration. Benoit-Vical et al.
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves Water, 80% methanol In vitro ND Cytotoxicity
Methanol extracts had minimal cytotoxicity with the aqueous extracts
exhibiting no cytotoxic effect.
Mudzwiri (2007)
Ceratotheca triloba
(Bernh.) Hook.f.
Leaves, roots Methanol, water In vitro ND Cytotoxicity
Puried extracts of Ceratotheca triloba had 41.473.2% HepG
cell death at
12.5 mg/25 ml while low concentrations (1.56 mg/ 25 ml) exhibited 114.4%
cell death.
Mohanlall (2010)
ND Not dened.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209221 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 puried 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 owering, 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 Olontoye (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.
10. Conclusions
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 mans 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 diversication, improved nutrition and human
health. However, more work is still needed to further validate the
vegetables 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 benets
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 (avonoids, 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 plantsbiologicalactivitiesinin
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 signicance 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
high quantities.
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
the manuscript.
Abubakar, M.S., Musa, A.M., Ahmeda, A., Hussaini, I.M., 20 07. The perception and
practice of traditional medicine in the treatment of cancers and inammations
by the Hausa and Fulani tribes of Northern Nigeria. J. Ethnopharmacol. 111,
Achigan-Dako, E., NDanikou, S., Assogba-Komlan, F., Ambrose-Oji, B., Ahanchede,
A., Pasquini, M., 2011. Diversity, geographical, and consumption patterns of
traditional vegetables in sociolinguistic communities in Benin: implications for
domestication and utilization. Econ. Bot. 65, 129145 .
Adéoti, K., Dansi, A., Ahoton, L., Kpèki, B., Ahohuendo, B., Ahanchédé, A., Vodouhè,
R., Hounhouigan, J., Sanni, A., 2009. Selection of sites for the in situ conserva-
tion of four traditional leafy vegetables consumed in Benin. Int. J. Biol. Chem.
Sci. 3, 13571374.
Aguilera, E., Lassaletta, L., Sanz-Cobena, A., Garnier, J., Vallejo, A., 2013. The potential
of organic fertilizers and water management to reduce N
O emissions in Medi-
terranean climate cropping systems. A review. Agric. Ecosyst. Env. 164, 3252.
Akula, U.S., Odhav, B., 2008. In vitro 5-Lipoxygenase inhibition of polyphenolic
antioxidants from undomesticated plants of South Africa. J. Med. Plants Res. 2,
Aletor, O., Oshodi, A.A., Ipinmoroti, K., 2002. Chemical composition of common
leafy vegetables and functional properties of their leaf protein concentrates.
Food Chem. 78, 6368.
Ayantunde, A.A., Hiernaux, P., Briejer, M., Udo, H., Tabo, R., 2009. Uses of local plant
species by agropastoralists in south-western Niger. Ethnobot. Res. Appl. 7,
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209221220
Barminas, J.T., Charles, M., Emmanuel, D., 1998. Mineral composition of non-con-
ventional leafy vegetables. Plant Foods Hum. Nutr. 53, 2936.
Bedigian, D., 2003. Evolution of sesame revisited: Domestication, diversity and
prospects. Genet. Resour. Crop. Evol. 50, 779787.
Bedigian, D., 2004. Slimy leaves and oily seeds: distribution and use of wild re-
latives of sesame in Africa. Econ. Bot. 58, S3S33.
Bedigian, D., Adetula, A.O., 2004. Ceratotheca sesamoides Endl. Prota 2: Vegetables/
Legumes. PROTA, Wageningen, The Netherlands.
Benoit-Vical, F., Soh, P.N., Saléry, M., Harguem, L., Poupat, C., Nongonierma, R., 2008.
Evaluation of Senegalese plants used in malaria treatment: focus on Chrozo-
phora senegalensis. J. Ethnopharmacol. 116, 4348.
Bolton, E.R., 1919. Note on the oil of Ceratotheca sesamoids. Analyst 44, 233234.
Dansi, A., Adjatin, A., Adoukonou-Sagbadja, H., Akpagana, K., 2008. Production and
traditional seed conservation of leafy vegetables in Benin rural areas. Bull. Rech.
Agron. Bénin 59, 5969.
Dansi, A., Vodouhè, R., Azokpota, P., Yedomonhan, H., Assogba, P., Adjatin, A., Loko,
Y., Dossou-Aminon, I., Akpagana, K., 2012. Diversity of the neglected and un-
derutilized crop species of importance in Benin, Sci. World J.
Diarra, N., Klooster, C., Togola, A., Diallo, D., Willcox, M., Jong, J., 2015. Ethnobota-
nical study of plants used against malaria in Sélingué subdistrict, Mali. J. Eth-
nopharmacol. 166, 352360.
Dobrian, A.D., Lieb, D.C., Cole, B.K., Taylor-Fishwick, D.A., Chakrabarti, S.K., Nadler, J.
L., 2011. Functional and pathological roles of the 12- and 15-lipoxygenases.
Prog. Lipid Res. 50, 115131.
Duncan, G., 2011. 715. Ceratotheca triloba. Curtis's Bot. Mag. 28, 205212.
Fasakin, K., 2004. Proximate composition of bungu (Ceratotheca sesamoides Endl.)
leaves and seeds. Biokemistri 16, 8892.
Fasakin, K., Olontoye, J.A., 2005. Agronomic characteristics of Bungu (Cerototheca
sesamoides Endl.). II: Effects of row spacing and seeding rate on leaf yield and
yield components. J. Agric. Res. Dev. 4, 5869.
Fasola, T., Ogunsola, O., 2014. The proximate and phytochemical composition of
Sesamum indicum Linn and Ceratotheca sesamoides Endl at different stages of
growth. J. Biol. Agric. Healthc. 4, 8488.
Grubben, G.J.H., Denton, O.A., 2004. Ceratotheca sesamoides Endl. In: Bedigian, D.,
Adetula, O.A. (Eds.), Plant Resources of Tropical Africa, vol. 2. VegetablesPROTA
Foundation. Backhuys Publishers, Wageningen, Netherlands.
Gülçin, İ., 2012. Antioxidant activity of food constituents: an overview. Arch. Tox-
icol. 86, 345391.
Heyman, H.M., Hussein, A.A., Meyer, J.J.M., Lall, N., 2009. Antibacterial activity of
South African medicinal plants against methicillin resistant Staphylococcus
aureus. Pharm. Biol. 47, 6771.
Hurrell, R.F., 2003. Inuence of vegetable protein sources on trace element and
mineral bioavailability. J. Nutr. 133, 2973S2977S.
Hurrell, R.F., Juillerat, M.A., Reddy, M.B., Lynch, S.R., Dassenko, S.A., Cook, J.D., 1992.
Soy protein, phytate, and iron absorption in humans. Am. J. Clin. Nutr. 56,
Hutchings, A., Scott, A.H., Lewis, G., Cunningham, A., 1996. Zulu medicinal plants:
an inventory. University of Natal Press, Pietermaritzburg, South Africa.
Ihlenfeldt, H.D., 2004. Pedaliaceae. In: Kadereit, J.W. (Ed.), Flowering Plants Dico-
tyledons. Springer, Berlin, Heidelberg.
Jäger, A.K., Hutchings, A., Van Staden, J., 1996. Screening of Zulu medicinal plants
for prostaglandin-synthesis inhibitors. J. Ethnopharmacol. 52, 95100.
Jeng, K.C.G., Hou, R.C.W., 2005. Sesamin and sesamolin: nature therapeutic lignans.
Curr. Enzym. Inhib. 1, 1120.
Konan, Y., Witabouna, K.M., Bassirou, B., Kagoyire, K., 2014. Antioxidant activity and
total phenolic content of nine plants from Côte dIvoire (West Africa). J. Appl.
Pharm. Sci. 4, 036041.
Lykke, A.M., Mertz, O., Ganaba, S., 2002. Food consumption in rural Burkina Faso.
Ecol. Food Nutr. 41, 119153.
Maharaj, R., Maharaj, V., Newmarch, M., Crouch, N.R., Bhagwandin, N., Folb, P.I.,
Pillay, P., Gayaram, R., 2010. Evaluation of selected South African ethnomedic-
inal plants as mosquito repellents against the Anopheles arabiensis mosquito in
a rodent model. Malar. J. 9, 301.
Maggioni, L., 2004. Conservation and use of vegetable genetic resources: a Eur-
opean perspective. Acta Hortic. 637, 1330.
Maroyi, A., 2011. Potential role of traditional vegetables in household food security:
a case study from Zimbabwe. Afr. J. Agric. Res. 6, 57205728.
McGaw, L.J., Jäger, A.K., Van Staden, J., 2000. Antibacterial, anthelmintic and anti-
amoebic activity in South African medicinal plants. J. Ethnopharmacol. 72,
Mertz, O., Lykke, A., Reenberg, A., 2001. Importance and seasonality of vegetable
consumption and marketing in Burkina Faso. Econ. Bot. 55, 276289.
Mitchikpe, E.C.S., Dossa, R.A.M., Ategbo, E.-A.D., van Raaij, J.M.A., Hulshof, P.J.M.,
Kok, F.J., 2008. The supply of bioavailable iron and zinc may be affected by
phytate in Beninese children. J. Food Comp. Anal. 21, 1725.
Mohanlall, V., 2010. In-vitro Propagation, Bioaugmentation and Screening of Cer-
atoteca triloba for the Production of Secondary Metabolites. Department of
Biotechnology and Food Technology, Durban University of Technology, Durban,
South Africa.
Mohanlall, V., Odhav, B., 2013. Antibacterial, anti-inammatory and antioxidant
activities of anthraquinones from Ceratotheca triloba (Bernh) Hook F. J. Med.
Plants Res. 14, 877886.
Mohanlall, V., Steenkamp, P., Odhav, B., 2011. Isolation and characterization of
anthraquinone derivatives from Ceratotheca triloba (Bernh.) Hook F. J. Med.
Plants Res. 5, 31323141.
Molander, M., Nielsen, L., Søgaard, S., Staerk, D., Rønsted, N., Diallo, D., Chifundera,
K.Z., Van Staden, J., Jäger, A.K., 2014. Hyaluronidase, phospholipase A
protease inhibitory activity of plants used in traditional treatment of snakebite-
induced tissue necrosis in Mali, DR Congo and South Africa. J. Ethnopharmacol.
157, 171180.
Moon, J.-K., Shibamoto, T., 2009. Antioxidant assays for plant and food components.
J. Agric. Food Chem. 57, 16551666 .
Mudzwiri, M., 2007. Evaluation of Traditional South African Leafy Plants for Their
Safety in Human Consumption. Department of Biotechnology, Durban Uni-
versity of Technology, Durban, South Africa.
NDanikou, S., Achigan-Dako, E., Wong, J.G., 2011. Eliciting local values of wild ed-
ible plants in southern Bénin to identify priority species for conservation. Econ.
Bot. 65, 381395.
Nadembega, P., Boussim, J.I., Nikiema, J.B., Poli, F., Antognoni, F., 2011. Medicinal
plants in Baskoure, Kourittenga Province, Burkina Faso: an ethnobotanical
study. J. Ethnopharmacol. 133, 378395.
Nura, S., Adamu, A.K., Mu'Azu, S., Dangora, D.B., Shehu, K., Mahamud, F.M., Mansur,
K., 2014. Assessment of the growth responses of sesame (Sesamum indicum l.)
and false sesame (Ceratotheca sesamoides Endl.) to colchicine treatments. Am. J.
Exp. Agric. 4, 902912.
Nura, S., Adamu, A.K., MuAzu, S., Dangora, D.B., 2012. Mutagenic effects of col-
chicine on some quantitative traits of false sesame (Ceratotheca sesamoides
Endl.). Savannah J. Agric. 7, 3945.
Obi, R.K., Iroagba, I.I., Ojiako, O.A., 2006. Virucidal potential of some edible Nigerian
vegetables. Afr. J. Biotechnol. 5, 17851788.
Odhav, B., Beekrum, S., Akula, U., Baijnath, H., 2007. Preliminary assessment of
nutritional value of traditional leafy vegetables in KwaZulu-Natal, South Africa.
J. Food Comp. Anal. 20, 430435.
Odhav, B., Kandasamy, T., Khumalo, N., Baijnath, H., 2010. Screening of African
traditional vegetables for their alpha-amylase inhibitory effect. J. Med. Plants
Res. 4, 15021507.
Ohkawa, A., Sakai, T., Ohyama, K., Fujimoto, Y., 2012. Malonylated glycerolipids from
the glandular trichome exudate of Ceratotheca triloba. Chem. Biodiv. 9,
Pickering, H., Roe, E., 2009. Wild owers of Victoria Falls Area Helen Pickering,
Pooley, E., 1998. A Field Guide to Wild Flowers KwaZulu-Natal and Eastern Region.
Flora Publications Trust, Durban, South Africa.
Rådmark, O., Samuelsson, B., 2007. 5-Lipoxygenase: regulation and possible in-
volvement in atherosclerosis. Prostaglandins Other Lipid Mediat. 83, 162174.
Raghuvanshi, R.S., Singh, R., 2001. Nutritional composition of uncommon foods and
their role in meeting in micronutrient needs. Int. J. Food Sci. Nutr. 32, 331335.
Raimondo, D., Von Staden, L., Foden, W., Victor, J.E., Helme, N.A., Turner, R.C., Ka-
mundi, D.A., Manyama, P.A., 2009. Red List of South African Plants 2009, Stre-
litzia 25. South African National Biodiversity Institute (SANBI), Pretoria, South
Roberts, M., 1990. Indigenous Healing Plants. Creda Press, Cape Town, South Africa.
Seifried, H.E., Anderson, D.E., Fisher, E.I., Milner, J.A., 20 07. A review of the inter-
action among dietary antioxidants and reactive oxygen species. J. Nutr. Bio-
chem. 18, 567579.
Surh, Y.-J., 2002. Anti-tumor promoting potential of selected spice ingredients with
antioxidative and anti-inammatory activities: a short review. Food Chem.
Toxicol. 40, 10911097.
Tarus, P.K., Machocho, A.K., Langat, T.C.C., Chhabra, S.C., 2002. Flavonoids from Te-
phrosia aequilata. Phytochemistry 60, 375379.
Toyin, M.Y., Khadijat, O.F., Saoban, S.S., Olakunle, A.T., Abraham, B.F., Luqman, Q.A.,
2012. Antidiarrhoeal activity of aqueous leaf extract of Ceratotheca sesamoides
in rats. Bangl. J. Pharmacol. 7, 1420.
Tredgold, M.H., 1986. Food Plants of Zimbabwe. Mambo Press, Gweru, Zimbabwe.
Uddin, I., Adesiyun, A., 2011. Insects associated with Bungu, Ceratotheca sesamoides
Endl. (Pedaliaceae) Ilorin, Nigeria. Agric. Biol. J. N. Am. 2, 974980.
Uddin II, R., Abdulazeez, R., 2013. Comparative efcacy of neem (Azadirachta indica),
false sesame (Ceratotheca sesamoides) Endl. and the physic nut (Jatropha curcas)
in the protection of stored cowpea (Vigna unguiculata) L. Walp. against the seed
beetle Callosobruchus maculatus (F.). Ethiop. J. Environ. Stud. Manag. 6,
Uddin II, R.O., Abdulazeez, R.W., 2013. Comparative efcacy of neem (Azadirachta
indica), false sesame (Ceratotheca sesamoides) Endl. and the physic nut (Jatropha
curcas) in the protection of stored cowpea (Vigna unguiculata) L. Walp. against
the seed beetle Callosobruchus maculatus (F.). Ethiop. J. Environ. Stud. Manag. 6,
Uusiku, N.P., Oelofse, A., Duodu, K.G., Bester, M.J., Faber, M., 2010. Nutritional value
of leafy vegetables of sub-Saharan Africa and their potential contribution to
human health: A review. J. Food Compos. Anal. 23, 499509.
Van Wyk, B.-E., Gericke, N., 2000. Peoples Plants: A Guide to Useful Plants of
Southern Africa. Briza Publications, Pretoria, South Africa.
Vanderjagt, D.J., Freiberger, C., Vu, H.T.N., Mounkaila, G., Glew, R.S., Glew, R.H.,
2000. The trypsin inhibitor content of 61 wild edible plant foods of Niger. Plant
Foods Hum. Nutr. 55, 335346.
Venter, C.S., Van Eyssen, E., 2001. More legumes for better overall health. South Afr.
J. Clin. Nutr. 14, S32S38.
Watt, J., Breyer-Brandwijk, M., 1962. The Medicinal and Poisonous Plants of
Southern and Eastern Africa. Livingstone, London, United Kingdom.
Yadang, G., Tchatchueng, J.-B., Tchiegang, C., 2009. Protein, carbohydrate, fat and
energy content of ‘‘ready-to-eat foods’’ in Cameroonian Sahels Region. J. Food
Technol. 7, 14.
N.A. Masondo et al. / Journal of Ethnopharmacology 178 (2016) 209221 221
... Frontiers in Agronomy | widely agreed that NUS are high in macro and micronutrients, in particular, potassium, calcium, magnesium, and zinc, and vitamins such as K, C, E, and folate, essential oils and bioactive compounds such as flavonoids, antioxidants, phenolics, and carotenoids (Johns and Eyzaguirre, 2006;Liu et al., 2011;Hughes and Ebert, 2013;Nyadanu and Lowor, 2014;Masondo et al., 2016;Tele Ayenan and Ezin, 2016;Nitcheu Ngemakwe et al., 2017;Campanaro et al., 2019), all of which are important for nutrition security and human and livestock health (see Table 1). Nyamupingidza et al. (2013) noted that the nutritional and chemical diversity of NUS makes them ideal nutritional supplements in diets of people with compromised immune systems (e.g., HIV/AIDS), reduces the incidence of non-communicable diseases (NCDs), improves wellbeing, and increases longevity ( Table 1). ...
... Nyamupingidza et al. (2013) noted that the nutritional and chemical diversity of NUS makes them ideal nutritional supplements in diets of people with compromised immune systems (e.g., HIV/AIDS), reduces the incidence of non-communicable diseases (NCDs), improves wellbeing, and increases longevity ( Table 1). This is because they are high in macronutrients such as carbohydrates, proteins, fats, and inorganic constituents and have anti-diarrhoeal, anti-diabetic, antiplasmodial, antibacterial, and antiviral properties (Masondo et al., 2016). For example, Kersting's groundnut (Macrotyloma geocarpum) reduces the occurrence of atherosclerosis and coronary heart disease, as it has a low fat (1.0%) and sodium content (5.67 mg g −1 ; Ayenan and Ezin, 2016).These health benefits provided by NUS are inexpensive and readily available, especially for the poor. ...
... Despite the numerous favorable properties of NUS, the research revealed that they have a few negative attributes. The leaves of false sesame (Ceratotheca sesamoides) are bitter and have a laxative effect, prompting consumers to find ways of preparation to reduce these effects (Masondo et al., 2016). These preparation methods include the addition of onions and tomatoes to soften and reduce the bitterness of C. sesamoides' leaves or serve with other vegetables to reduce the laxative effect (Masondo et al., 2016). ...
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There are growing calls to adopt more sustainable forms of agriculture that balance the need to increase production with environmental, human health, and wellbeing concerns. Part of this conversation has included a debate on promoting and mainstreaming neglected and underutilized crop species (NUS) because they represent a more ecologically friendly type of agriculture. We conducted a systematic review to determine the ecosystem services derived from NUS and assess their potential to promote functional ecological diversity, food and nutritional security, and transition to more equitable, inclusive, sustainable and resilient agricultural landscapes and food systems in Africa. Our literature search yielded 35 articles for further analysis. The review showed that NUS provide various provisioning, regulating, cultural, and supporting ecosystem services and several environmental and health co-benefits, dietary diversity, income, sustainable livelihood outcomes, and economic empowerment, especially for women. Importantly, NUS address the three pillars of sustainable development- ecological, social, and economic. Thus, NUS may provide a sustainable, fit-for-purpose transformative ecosystem-based adaptation solution for Africa to transition to more sustainable, healthy, equitable, and resilient agricultural landscapes and food systems.
... (K, Hooker herbarium), Senegal, June 1821, states, "médicament des l'ombu des buissons" [bush medicine]. Numerous authors praise the nutritious leafy greens of Ceratotheca and Sesamum (e.g., Auwalu and Babatunde 2007;Masondo et al. 2016;Orech et al. 2005; Vihotogbe-Sossa et al. 2012), but exhaustive searches retrieved few studies about the micronutrient contents of the leaves. Reports that exist are difficult to compare because data are incomplete and inconsistent: (1) piecemeal investigations of micronutrients, (2) values given in tables differ from the authors' summaries of the data, and (3) results varied widely. ...
... Few studies have documented the usage (nutrition and pharmacology) of Ceratotheca sesamoides and C. triloba leaves (Masondo et al. 2016), although Morton (1990:256) and Bedigian (2013a) presented several therapeutic uses of their mucilaginous leaves as poultices on wounds, boiled or steeped in cold water as a remedy for chronic diarrhea, and other intestinal and urinary disorders, for ophthalmic and skin complaints, and as an allpurpose demulcent. ...
The usage and cultural importance of wild and weedy edible leaves of wild relatives of sesame, Ceratotheca and Sesamum in Africa, is reported from herbarium records, published sources, and firsthand observations. They contribute not only nutritionally-protein, lipid, and micronutrients, especially calcium and iron-but release a little scrutinized valuable product, water-soluble polysaccharide mucilage, that serves as a source of fiber and provides beneficial dietary bulk, enhances gastrointestinal function, alleviates constipation, protects against diabetes, and lowers cholesterol. Dissimilar genera and species are used in similar ways. Their utility was discovered by women gathering edible leaves to accompany dry cereal staples. African vernacular names from several disparate cultures refer to their slimy consistency. Corresponding nutritional and mucilage constituents of several other popular African vegetables are evaluated for comparison.
... Ceratotheca sesamoides is endemic to Africa and is present in most sub-Saharan African countries. It is locally cultivated as a legume crop for its leaves that have high nutrient properties (Masondo et al. 2016), and it is considered as a medicinal plant for its antidiarrheal and antiseptic properties (Fasola & Ogunsola 2014). Sesamum radiatum and C. sesamoides are very close species despite some differences in their phytochemical composition (Fasola & Ogunsola 2014) and both are called false sesame. ...
Strong full-plant watering and insecticide sprays were unfavorable for Nesidiocoris tenuis. • Ceratotheca sesamoides and Physalis micrantha were natural host plants of N. tenuis in Senegal. • Sesamum radiatum supported good N. tenuis development and reproduction without alternative food. • Olfactometry showed that N. tenuis would not abandon tomato crops for S. radiatum. A R T I C L E I N F O Keywords: Tuta absoluta Lycopersicon esculentum Insectarium plant Companion plant Oflactometry Sesamum radiatum A B S T R A C T Conservation biological control is a promising approach to natural pest control in both open field and protected crops. In sub-Saharan Africa, tomato crops are commonly damaged by several arthropod pests but can also host the predatory mirid bug, Nesidiocoris tenuis. Unfortunately, farmers do not currently take full advantage of this zoophytophagous and generalist biocontrol agent. We conducted field investigations in the Niayes area of Senegal, combined with laboratory experiments, to determine how to enhance natural regulation services provided by this natural enemy. We found that strong full-plant watering and insecticidal sprays, both conventionally applied in the Niayes area, had a detrimental effect on N. tenuis. We identified Sesamum radiatum (false or black sesame) as a promising insectarium plant that could also serve as a leaf vegetable and medicinal plant. This plant supports good N. tenuis development and reproduction, even without alternative food sources. Y-tube olfactometry indicated no attraction to odors of Sesamum radiatum, suggesting that N. tenuis would not abandon tomato plants for this insectarium plants when present in tomato fields. Our results suggest potential approaches to improve conservation of N. tenuis for biological control of Tuta absoluta and other tomato pests in Africa.
... Although it is plausible that early herbal healing practices developed in Africa, considered the cradle of humanity, there has remained relatively little systematic documentation, as this knowledge has mainly been passed through oral tradition (Okpako 1999). However, several recent publications show that the application of food as medicine is still widespread in different parts of Africa (Grimaldi & van Andel 2018, Kimondo et al. 2015, Masondo et al. 2016, Omokhua-Uyi & Van Staden 2020, Roulette et al. 2018, Towns & van Andel 2016 (Tan et al. 2017, Wu & Liang 2018, Zhang & Wang 2014, Zhao et al. 2021, Zou 2016). In addition, some references to handbooks on practical applications are provided at the end of this review for further reading. ...
Traditional foods and diets can provide health benefits beyond their nutrient composition because of the presence of bioactive compounds. In various traditional healthcare systems, diet-based approaches have always played an important role, which has often survived until today. Therefore, investigating traditional foods aimed at health promotion could render not only novel bioactive substances but also mechanistic insights. However, compared to pharmacologically focused research on natural products, investigating such nutrition-based interventions is even more complicated owing to interacting compounds, less potent and relatively subtle effects, the food matrix, and variations in composition and intake. At the same time, technical advances in ‘omics’ technologies, cheminformatics, and big data analysis create new opportunities, further strengthened by increasing insights into the biology of health and homeostatic resilience. These are to be combined with state-of-the-art ethnobotanical research, which is key to obtaining reliable and reproducible data. Unfortunately, socioeconomic developments and climate change threaten traditional use and knowledge as well as biodiversity. Expected final online publication date for the Annual Review of Food Science and Technology, Volume 13 is March 2022. Please see for revised estimates.
... Traditional African leafy vegetables are increasingly being recognized for their roles in nutrition (Abukutsa-Onyango, 2007;Jinazali et al., 2017;Masondo et al., 2016;Neugart et al., 2017), food supply (Almekinders and De Boef, 2000;Baldermann et al., 2016;Hughes and Ebert, 2013), ethno-botanical properties (Mokganya and Tshisikhwa, 2019;Moteetee et al., 2019) and plant breeding (Cernansky, 2015;Kiebre et al., 2015;Nyadanu et al., 2017;Wasonga et al., 2015). They are easily accessible, with minerals' and vitamins' levels exceeding those found in most exotic vegetables (Abukutsa-Onyango, 2007;Nyadanu and Lowor, 2015). ...
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Traditional leafy vegetables are increasingly being recognized as parents in plant breeding, and source of improved nutrition and food supply. Wild cucumber (Cucumis myriocarpus), a traditional leafy vegetable has been reported to possess some resistance to root-knot (Meloidogyne species) nematodes; however, the form of nematode resistance in these plants is not documented. The objective of this study was to determine the possible mechanism of nematode resistance in C. myriocarpus to M. incognita and M. javanica under greenhouse conditions. Two separate experiments of C. myriocarpus were each inoculated with second-stage juveniles (J2) of M. incognita or M. javanica. Harvesting of seedlings was conducted every other day from 2 days after inoculation. At each harvest, roots of five seedlings were stained using acid fuchsin method for assessment and number of necrotic spots, giant cells, rootlet interference, root galls and nematode juveniles counted. Harvest period had highly significant effects (P ≤ 0.01) on numbers of giant cells, rootlet interference and root galls, but were not significant on necrotic spot number in C. myriocarpus - M. javanica relations. Harvest interval were significant (P ≤ 0.05) on giant cell number and highly significant on root gall number in C. myriocarpus - M. incognita relations. The assessment of nematode infection showed that there was delayed penetration of both M. incognita and M. javanica in C. myriocarpus but their ability to cause disease was impaired. In conclusion, post-infectional resistance is the possible mechanism in C. myriocarpus traditional leafy vegetable against the two Meloidogyne species.
... In terms of nutritional value, the plant has high energy value and fat content (Odhav et al. 2007). Furthermore, the leafy vegetable is widely utilized for its medicinal properties for certain maladies including painful menstruation, stomach cramps, nausea, fever, and diarrhea which have been vastly attributed to the diverse phytochemicals in the plant (Masondo et al. 2016a). Despite the uses and value of C. triloba, there is still a lack of research on its cultivation practices as the plants are mostly found in the wild. ...
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We evaluated the role of Kelpak® on growth, nutritional content, phytohormones, and phenolic acids in the angiosperm Ceratotheca triloba (Bernh.) Hook.f. grown under macronutrient-deficient conditions. Seeds were germinated in the greenhouse and established seedlings were treated with 50% Hoagland’s solution (50% HS) or nutrient-deficient solutions lacking one of the following: nitrogen (-N), phosphorus (-P), potassium (-K), which were either treated with or without Kelpak® (control). After 4 months of growth, macronutrient deficiency severely reduced growth and nutritional composition in plants. Nevertheless, the detrimental effects of insufficient macronutrient supply on growth were ameliorated with the application of Kelpak®, with a fourfold (-N), threefold (-P), and twofold (-K) increase in leaf and root fresh weight. Mineral and carbohydrate levels increased in plants deprived of N and P when Kelpak® was applied in the different soils. Macronutrient stress (N, P, K) increased phytohormone production in C. triloba more than in 50% HS-treated plants. The phytohormone content in macronutrient-deficient leaves was further enhanced with the application of Kelpak®. On the contrary, application of Kelpak® suppressed the production of phenolic acids in 50% HS, -N, and -K-treated plants when compared to the control treatments. The interaction between macronutrient deficiency and Kelpak® application played an important role in the regulation of phytohormone and phenolic acid concentration in C. triloba. The current findings demonstrated the potential of Kelpak® as an efficient biostimulant that enhances nutrient uptake for improved growth and biochemicals in C. triloba under macronutrient stress condition.
... The species serves as a supplement to the starch-based diets of under-developed communities in southern Africa, and its nutritional content has been documented (Odhav et al. 2007). The extensive biological activities of C. triloba have been mainly attributed to phytochemicals such as phenolics, steroids, and alkaloids commonly present in the plant (Masondo et al. 2016a). These include several in vitro and ex vitro studies reported on the antioxidants, antimicrobial, anti-diabetic, and antiplasmodial properties of the plant. ...
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We evaluated the effect of different watering regimes on the growth, chlorophyll fluorescence, phytohormones, and phenolic acids in Ceratotheca triloba (Bernh.) Hook.f., a commonly consumed African indigenous leafy vegetable. The study was conducted in the greenhouse under different watering regimes [seven (daily); three (thrice); two (twice); one (once) day(s) per week] for a period of 2 and 4-months. In each pot (7.5 cm diameter; 150 ml volume), 50 ml of water was applied per treatment. At the end of the experiment, plant growth, chlorophyll fluorescence, phytohormones, and phenolic acids were determined. A decrease in water availability resulted in a consistent decline in plant growth after a 4-month growth period. The severity of reduced water availability was more noticeable in plants watered once a week with a 1.4-fold reduction in growth and quantum efficiency of PSII (Fv/Fm) value of 0.80. The significant decline in growth and chlorophyll fluorescence was probably due to the increased production of abscisic acid (ABA) and cytokinin (CK) content together with the detected phytohormones in plants with restricted water supply. Furthermore, plants watered once a week had a trade-off between growth and phenolic acid production, with significantly higher (threefolds) concentrations of vanillic, ferulic, caffeic, and 4-coumaric acids in 4-month-old plants. Even though C. triloba grew best in well-watered soil, the plant had the potential to adapt and survive in soils with limited water supply for longer periods of growth. These findings suggest that regulation of phytohormones and phenolic acids played an important role in improving the growth of C. triloba under limited water conditions.
... In response to these changes, plants improve metabolic profile or generate different secondary metabolites (Chen and Arora, 2011). These stress treatments also improve the growth, germination, abiotic and salt tolerance of vegetables (Masondo et al., 2016). The agriculturists, farmers and food scientists are trying to explore more sustainable and viable approach to enhance phenolic attributes (secondary metabolites) without compromising the crop yield (Duan et al., 2006). ...
The current study focuses on to the latent potential of Plant Leaf Extract (PLE) to induce growth and antioxidant character in radish root when applied as pre-sowing seed priming agents. The growth behavior in primed seed with PLE of mulberry leaf (mL), Brassica leaf (BL), Sorghum leaf (SL), Moringa leaf (MoL), was regularly monitored whereas, proximate composition, biochemical attributes and antioxidant potential was assessed after 60 days of emergence. Results revealed that PLE based seed priming affect growth physiology consequently improved total phenolic content (TPC), radical scavenging capacity (DPPH IC50) and reducing power of radish root. Overall, seed bio-priming with 3% of MoL for 6 h was found to be the most effective to optimize growth with improved biochemical and antioxidant attributes. So far, seed bio-priming with crude extracts of selected plant leaves may provide viable bioactive fortification choice.
... (Pedaliaceae) is an indigenous leafy vegetable found in southern Africa. The nutritional, medicinal and pharmacological properties of the vegetable are well-documented (Masondo et al., 2016). Ceratotheca triloba is propagated through seed germination in spring or early summer in rich and well-drained soils. ...
Extreme temperatures, drought and salinity stress adversely affect seed germination and seedling growth in crop species. Seed priming has been recognized as an indispensable technique in the production of stress-tolerant plants. Seed priming increases seed water content, improves protein synthesis using mRNA and DNA and repair mitochondria in seeds prior to germination. The current study aimed to determine the role of biostimulants-seed-priming during germination and seedling growth of Ceratotheca triloba (Bernh.) Hook.f. (an indigenous African leafy vegetable) under low temperature, low osmotic potential and salinity stress conditions. Ceratotheca triloba seeds were primed with biostimulants [smoke-water (SW), synthesized smoke-compound karrikinolide (KAR1), Kelpak(®) (commercial seaweed extract), phloroglucinol (PG) and distilled water (control)] for 48h at 25°C. Thereafter, primed seeds were germinated at low temperatures, low osmotic potential and high NaCl concentrations. Low temperature (10°C) completely inhibited seed germination. However, temperature shift to 15°C improved germination. Smoke-water and KAR1 enhanced seed germination with SW improving seedling growth under different stress conditions. Furthermore, priming seeds with Kelpak(®) stimulated percentage germination, while PG and the control treatment improved seedling growth at different PEG and NaCl concentrations. Generally, high concentrations of PEG and NaCl brought about detrimental effects on seed germination and seedling growth. Findings from this study show the potential role of seed priming with biostimulants in the alleviation of abiotic stress conditions during seed germination and seedling growth in C. triloba plants.
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Using participatory research appraisal 29 villages randomly selected in different ethnic areas of Benin were surveyed to document the traditional practices and local knowledge pertaining to the production and the seed system of the traditional leafy vegetables (TLVs) in Benin. The results revealed that in rural areas of Benin, TLVs are cultivated at a subsistence level with occasionally use of chemicals providing hence a good opportunity for the promotion of organic farming. The seed system and on farm conservation are still archaic and should be improved. In the north and during the dry season, leafy vegetables are traditionally dried under direct sunlight for post-harvest storage. The need to improve the drying methods by investigating the use of solar dryers with a view of minimizing nutrient loss and increasing the amount of vegetables preserved was highlighted. Development of awareness of the merits of the TLVs, promotion of seeds marketing, development of agronomic and utilisation packages (including pest and diseases control) on some species (Acmella oleracea, Ceratotheca sesamoides, Crassocephalum rubens, Justicia tenella, Launaea taraxacifolia, Sesamum radiatum and Solanum scabrum), selection and promotion of high yielding varieties and financial assistance to organized farmer groups were key interventions suggested for the promotion of TLVs in Benin. The study highlighted that women play an important role in the production of the TLVs and led to the conclusion that in their traditional roles of housewives, plant gatherers, home gardeners, plant domesticators, herbalists and seed custodians they are mainly responsible to the preservation and the sustainable use of the TLVs and should be considered as key actors in any programme of TLVs conservation and valorisation in Benin.
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Sesamum indicum Linn. and Ceratotheca sesamoides Endl., both of family Pedaliaceae which were commonly harvested in substitute of one another now belong to the group of neglected and underutilized indigenous vegetables of the tropics. There is no known record of their chemical composition in line with their growth. In the light of this, the proximate and phytochemical composition of these closely related plants were studied in order to bring to limelight the potentials of these increasingly neglected vegetables. Significant differences were observed in the percentage proximate and phytochemical analyses of the two species at different stages of growth. Comparative studies also revealed a higher percentage proximate in S. indicum (9.57% crude fibre, 9.53% moisture) than C. sesamoides (8.67% crude fibre, 9.17% moisture) while the phytochemicals were significantly more in the S. indicum (138.33mg/100g Tannins) than C. sesamoides (127mg/100g Tannins). The values for the two species were at par for saponins at 330mg/100ml, while the other phytochemicals present had slightly higher values in C. sesamoides for flavonoids, alkaloids and phenols at 1315 mg/100g, 843 mg/100g and 46.3mg/100g respectively. In this study, proximate, mineral and phytochemical constituents showed a decline as the leaves approached reproductive stage of flowering and fruiting. It was noted in the two species studied that S. indicum started flowering after 8 weeks of growth and C. sesamoides, between the 9th and the 10th week. The proximate and the phytochemical values of the plants strongly support their use as vegetables and suggestive of medicinal values hence should not be neglected. Key words: Proximate analysis, Phytochemicals, Sesamum indicum and Ceratotheca sesamoides
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Extracts from four leaves: Ceratotheca sesamoides (Endl.), Lactuca taraxacifolia (Willd.), Euphorbia lateriflora (Schum. & Throes) and Amaranthus viridis (Linn.) used in Nigeria as edible vegetables were tested for antiviral activities using the measles virus on Human Epidermoid carcinoma (HEP-2) cell line. The efficacy of each extract in preventing viral penetration into a living cell as well as its effect on the virus itself and the toxicity of same on the cell were all determined. All the extracts showed dose-dependent antiviral activities. The inhibitory action of the extracts from C. sesamoides and L. taraxacifolia were greatest at high concentrations of 15 mg/ml when the virus had already penetrated the cell. E. lateriflora only inhibited the virus activity at very high concentrations of 15 mg/ml when the virus was treated with the extract and also when the HEP-2 cell was treated with the extract before the virus suspension was added. Amaranthus viridis concentrations of 10 and 15 mg/ml were contaminated but the extract at a low concentration of 5 mg/ml was able to exhibit antiviral activity when the cell was treated with the extract before the virus was added. In view of the efficacy of these extracts in inhibiting the cytopathic effect (CPE) of measles virus without harming the host, the utilization of the extracts in the formulation of new antiviral drugs is strongly recommended especially when the availability and the very low cost of these vegetables are considered.