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J. Bio. & Env. Sci. 2020
101 | Laizer et al.
REVIEW PAPER OPEN ACCESS
Insights of allelopathic, insecticidal and repellent potential of
an invasive plant Sphaeranthus suaveolens in pest and weed
management
Hudson Laizer*1,2
, Musa Chacha1
, Patrick Ndakidemi1
1
Department of Sustainable Agriculture and Biodiversity Conservation, Nelson Mandela African
Institution of Science and Technology, Arusha, Tanzania
2
Centre for Research, Agricultural Advancement, Teaching Excellence and Sustainability in Food
and Nutritional Security (CREATES), The Nelson Mandela African Institution of Science and
Technology (NM-AIST), Arusha, Tanzania
Article published on August 30, 2020
Key words: Botanical extracts, Secondary metabolites, Sustainable agriculture, Smallholder farmers, Terpene
Abstract
Sphaeranthus suaveolens is a weed from the family Asteraceae, it grows abundantly in wet areas and most
common in rice fields. The extracts from plants closely related to S. suaveolens have been reported to have
allelopathic, insecticidal, antifeedant, repellent, and other biological activities. Currently, the use of synthetic
chemicals to control weeds and insect pests raise several concerns related to environment and human health.
Extracts from plants with pesticidal properties can offer the best and an environmentally friendly alternative.
Some of these extracts have been extensively tested to assess their applications as valuable natural resources in
sustainable agriculture. This review article therefore explores the potential of S. suaveolens extracts in
controlling insect pests and managing weeds by smallholder farmers.
*Corresponding Author: Hudson Laizer  laizerh@nm-aist.ac.tz
Journal of Biodiversity and Environmental Sciences (JBES)
ISSN: 2220-6663 (Print) 2222-3045 (Online)
Vol. 17, No. 2, p. 101-112, 2020
http://www.innspub.net
J. Bio. & Env. Sci. 2020
102 | Laizer et al.
Introduction
Agricultural production in most part of the world is
affected by pests, diseases and weeds among other
factors (Nations, 2016; Singh et al., 2003). Majority
of the farmers respond to these constraints through
the use of synthetic chemicals such as herbicides and
pesticide (Kelly et al., 2003). The extensive use of
these chemicals in controlling insect pests and
managing weeds, however, have alarmed the public
on the effects they might bring to human health and
the environment at large (Khanh et al., 2005). Such
concerns are putting pressure on agricultural sector
to reduce the use of chemicals and as a result, much
attention is paid to alternative methods and
techniques for controlling and managing weeds and
insect pests, through non-chemical methods and/or
use of natural products such as botanical extracts
(Isman, 2006a; Williamson et al., 2008).
Prior to the discovery and commercialization of
synthetic pesticides, botanical extracts, among other
methods, were used by most farmers in crop
protection against insect pests, weeds and diseases
(El-Wakeil, 2013). Extracts from plants with
allelopathic or pesticidal properties were of great
importance in making natural herbicides and
pesticides (Benner, 1993; Godfrey, 1994). There is
also increasingly evidence from literatures that plant
extracts can be manipulated and used as perfect
agrochemicals in controlling insect pests and
managing weeds (Hoagland, 2001; Macías et al.,
2001; Mkenda et al., 2015; Nattudurai et al., 2012;
Ngondya et al., 2016; Singh et al., 2003; Stephen et
al., 2002; Vyvyan, 2002).
The secondary metabolites in plants are responsible
to biological activities that offer defense against
predators, fungi and bacteria, also these metabolites
may act as natural herbicides by suppressing other
plant species (Dewick, 2009; Schoonhoven et al.,
2005). These biological activities from the plants
secondary metabolites can be exploited and
manipulated for various human uses, and in this
respect Sphaeranthus suaveolens has a considerable
potential. S. suaveolens is a widespread weed in
swampy and irrigated farmlands, and usually infests
cultivated fields and reduces crop productivity
(Beentje, 2002). A heavy infestation of this weed
results in adverse effects on the growth and yield of
crops, particularly in rice fields (Fahmy, 1997).
It has been observed that S. suaveolens has an ability
to overcome and suppress crop plants in a wide range
over a short period of time (Ivens, 1989). However,
the secondary metabolites involved are largely
unknown and weather they can be applied in
managing other weeds and controlling insect pests is
yet to be determined. Understanding this could
considerably justify the practical application of
botanical-based weeds and insect pests management
techniques for most smallholder farmers in areas
where S. suaveolens is growing.
This review article therefore highlights the
allelopathic, insecticidal and repellent potential of an
invasive plant S. suaveolens with a focus on its
application in controlling insect pests and managing
weeds by smallholder farmers.
Overview of S. suaveolens and its spatial distribution
in Tanzania
S. suaveolens is an aromatic annual spreading herb
from the family Asteraceae with broad sessile leaves
covered with glandular hairs (Osman, 2011). The lower
stem often trails along the ground and roots at the
nodes with thread-like root, flowers are purple, in
compound heads ovoid in shape and borne on solitary
glandular peduncles with toothed wings (Fayed &
Mohamed, 1991).
The head as a whole is surrounded by several rows of
bracts, of which only the tips are visible when flowers
are fully open, and propagated by seeds which takes
about 10-12 days to germinate, and the seedlings attain
the height of 5.0 - 6.0cm within 30 days in a favorable
environment (Beentje, 2002).
S. suaveolens is widespread in Africa over a range of
altitudes from Rwanda, Burundi, Sudan, Ethiopia,
J. Bio. & Env. Sci. 2020
103 | Laizer et al.
Zambia, Malawi, Mozambique, Egypt, Uganda, Kenya
to Tanzania (Everard et al., 2002; Fahmy, 1997). It is
mostly found growing in wet areas and thrives well in
medium clayey soils, but also common in and around
irrigation ditches and rice fields and considered as a
major weed in most farms (Ivens, 1989). In Tanzania,
the plant was first reported in Songea (Brenan, 1960),
then Mpwapwa (Launert, 2003), Mkata and Mandela
in Wami River Ecosystem (Mligo, 2017). The spatial
distribution of S. suaveolens in Northern Tanzania
particularly Arusha and Kilimanjaro regions however,
have not been mapped despite being reported as a
weed to most agricultural fields.
Fig. 1. Distribution of S. suaveolens in Wami River Ecosystem in Tanzania.
Allelopathic effects of S. suaveolens on crops
Allelopathy is a phenomenon, whereby one plant
influences the growth of another one, including
microorganisms by the release of chemical
compounds into the environment (Keeley, 2010; Rice,
1983; Whittaker & Feeny, 1971). The allelopathic
effects are the result of chemical compounds known
as allelochemicals, which are usually plants’
secondary metabolites or byproducts of the principal
metabolic pathways in plants (Chancellor, 1987;
Dayan et al., 2009; Macías et al., 2007). In recent
years, allelopathy has become a research hotspot for
making comprehensive analysis about the mechanism
of weeds and identification of specific chemical
compounds responsible for allelopathic effects
(Azirak & Karaman, 2008; Bais et al., 2003; Einhellig
& Leather, 1988).
Most allelopathic plants have been observed to
significantly affect the growth, productivity and yield
of other crops by causing soil sickness and nutrient
imbalance (Kohli et al., 2008), as well as affecting the
microbial population (Batish et al., 2001). Several
studies have indicated that, most weeds possess
allelopathic effects which play a significant roles in
their invasion success (Macías et al., 2014; Qasem &
Foy, 2001; Zhou et al., 2013). Numerous weeds from
the Asteraceae family have been reported to possess
allelopathy and can significantly inhibit crop
productivity in agricultural land (Ilori et al., 2010;
Kong et al., 2007). The invasive weed, Sphaeranthus
indicus has been reported to inhibit seed germination
and growth of wheat (Triticum aestivum), rice (Oryza
J. Bio. & Env. Sci. 2020
104 | Laizer et al.
sativa) and mung bean (Vigna radiata) in different
farming systems (Lodha, 2004).
Recently, Mahajan et al. (2015) reviewed the
allelopathic potential of S. indicus and found that the
germination and seedling growth of various crops
were significantly decreased with increase in
concentration of its extract.
Despite of the negative effects on cultivated crops,
allelochemicals from allelopathic plants can be
manipulated and used to control weeds of various
crops. For example, Khanh et al. (2006) noted that
the allelochemicals contained in tissues of Passiflora
edulis can significantly suppress the two noxious
paddy rice weeds (Echinochloa crusgalli and
Monochoria vaginalis). Other studies have also
reported the use of allelochemicals for weed control in
the laboratory as well as application under field
conditions (Jabran et al., 2015; Ngondya et al., 2016).
Jamil et al. (2009) described the utilization of
allelopathic water extract as an important and useful
way of exploiting the allelopathic potential to manage
wild oat and canary grass in wheat fields.
The emergence and root length of most rice weeds
was inhibited by allelochemical (Lycorine) from the
dead leaves of spider lily (Iqbal et al., 2006). The
allelopathic crops when used as cover crop, mulch,
smother crops, green manures, or grown in rotational
sequences maybe helpful in reducing noxious weeds
and plant pathogen, improve soil quality and crop
yield (Khanh et al., 2005).
Furthermore, the application of allelopathic extracts
may give an efficient alternative control over weeds
similar to that offered by synthetic herbicides (Xuan
et al., 2004). Interactions among potential
allelopathic plants, target pests and other non- target
organisms in a cropping system also need to be
considered and fully realized to avoid detrimental
effects to desired crops and non-target species
(Farooq et al., 2013).
The allelochemicals involved in weed suppression can
serve as basic templates for developing new
generation of biopesticides with low or no toxic
effects to the environment and human health
(Ferguson et al., 2009).
The allelopathic potential of S. suaveolens, therefore
need to be realized and selectively used to suppress
density of other weeds and insect pests population
particularly in small farming agricultural systems.
Insecticidal and repellent activities of S. suaveolens
to insect pests
Insecticides, weather natural or synthetic are
developed to either kill, repel, or interfere with the
damaging behavior of insect pests (EPA, 2009). Due
to intensity of plant-insect interactions, plants have
well developed defense mechanisms against insect
pests by producing natural compounds which acts as
natural pesticides (Després et al., 2007). The most
exciting concept is to isolate and identify such
compounds and use them as candidates in making
safer pesticides (Maia & Moore, 2011). Plant extracts
with pesticidal properties can be active against
specific target insects, biodegradable, have low to
non-toxic effects, cheap and easy to prepare (Kim et
al., 2003; Mkindi et al., 2017). Due to this facts, these
plant extracts could lead to the development of new
classes of safer pesticides (Céspedes et al., 2014;
Tembo et al., 2018).
The Asteraceae family in which S. suaveolens belongs,
have been reported to contain plants with insecticidal
activities (Dhale, 2013; Green et al., 2017; Sosa et al.,
2018). These insecticidal activities are mostly linked
to presence of secondary metabolites such as
terpenes, which can act as larvicides, insect growth
regulators and feeding and oviposition deterrents
(Miresmailli & Isman, 2014). Terpene is among the
most diverse class of plant secondary metabolites
found in essential oils of most plants, S. suaveolens
included (Ahmed et al., 2017; Pagare et al., 2015).
This secondary metabolite has been reported to play
an important role in plant protection against
pathogens (Neerman, 2003), insects (Wu et al., 2016)
and toxic to mammals as well (Gurib-Fakim, 2006).
Recently, Sosa et al. (2018) reported insecticidal
activities of the terpene isolated from Vernonanthura
nebularum against fall army warm (Spodoptera
J. Bio. & Env. Sci. 2020
105 | Laizer et al.
frugiperda) and fruit fly (Ceratitis capitate).
Moreover, two other terpenes from Inula helenium
were examined by Kaur et al. (2017) and reported to
significantly inhibit the growth of tobacco leafworm
(Spodoptera litura).
Other techniques such as use of water extracts from S.
indicus was also reported to demonstrated toxic
effects against insect pests such as rice weevil (Patole
et al., 2008), cowpea weevil (Singh & Shrivastava,
2012) and red flour beetle (Pugazhvendan et al.,
2012). Furthermore, the extracts from S. indicus have
been showing larvicidal activities and repellent
activities to most of the insect pests (Arivoli et al.,
2016; Baby, 1994; Singh & Shrivastava, 2012).
The presence of secondary metabolites with pesticidal
properties such as terpene in the essential oil of S.
suaveolens may give positive insecticidal and
repellent activities to most insect pests, hence used
for as protective agents against insect pests, but this
needs further scientific investigation.
Key allelochemicals from the leaf, stem and root
extracts of S. suaveolens
Allelochemicals are secondary metabolites produced
by living organisms such as plants that have
stimulatory or inhibitory effects upon the growth,
health, behavior and distribution of neighboring
organisms being another plants, insects or microbes
(Haig, 2008). The role played by secondary
metabolites is mostly ecological, linked to plant
defense against other plants, pests, or diseases
(Ramakrishna & Ravishankar, 2011). Allelochemicals
undoubtedly pose problems in agriculture, but if well
manipulated they can be beneficial and offer great
opportunities such as insect pests and weeds control
(Einhellig, 1987). Despite the efforts in allelopathic
researches, little is known on the potential to exploit
the key allelochemicals in agricultural systems and
use them as templates in making safer and affordable
herbicides and/or pesticides (Kremer & Ben-
Hammouda, 2009). Much of the work to date has
focused on weather extracts from S. suaveolens show
biological activities such as antimicrobial, immune
stimulating, anticancer, antitumor, anthelmintic,
repellency, insecticidal and allelopathy (Ahmed &
Mahmoud, 1997; Kleinowski et al., 2016). However,
very few literatures have reported the identified
compounds found in S. suaveolens extracts, none of it
has a list of allelochemicals found in S. suaveolens
despite being the weed of economic importance in
many rice and common bean farms in Africa.
Allelochemicals belong to various chemical groups,
and can be classified based on their structures and
properties into: water-soluble organic acids, straight-
chain alcohols, aliphatic aldehydes, and ketones,
lactones, long-chain fatty acids and polyacetylenes,
quinines (benzoquinone, anthraquinone and complex
quinines), phenolics, cinnamic acid and its
derivatives, coumarins, flavonoids, tannins, steroids
and terpenoids (Li et al., 2010).
Most of these biochemicals are synthesized during the
shikimate pathway (Hussain & Reigosa, 2011) or, in
the case of essential oils, from the isoprenoid pathway
(Rehman et al., 2016). The extract of the aerial parts
of S. suavealens was reported by Jakupovic et al.
(1990) to contain eight eudesman-12. 6 β abides,
carvotacetone derivatives and a thymohydroquinone
glucopyranoside. Later on, Pooter et al. (1991)
reported extract of the same plant comprise of
thymohydroquinone dimethylether, a diacetylene
thiophene, inositol and myoinositol esters, and
several carvotanacetone derivatives, he went further to
examine the essential oil of S. suaveolens and noted
methyl chavicol, α-ionon,e, dcadinene and p-
methoxycinnamadehyde as major constituents, and a-
terpinene, citral, geraniol, geranyl acetate, βionone,
shaerene, indicusene and sphaeranthol as minor
constituents. Ahmed and Mahmoud (1997) examine
the extract of aerial parts of S. suaveolens and reported
three carvotacetone derivatives, together with four
monoteroene compounds. Later on, Hassanali et al.
(1998) reported cis-pinocamphone as the major
constituents (63.5%) of the leaf oil of S. sauveolens.
The details of these identified compounds are stated
J. Bio. & Env. Sci. 2020
106 | Laizer et al.
in Table 1, however, identifying alone isn’t sufficient
enough, rather gaining an understanding on which
among these compounds are allelochemicals and how
to use them in improving crop production though
managing weeds and controlling insect pests in
sustainable agriculture will be a big advantage.
Table 1. Identified compounds from leaf oil of S.
suaveolens.
Compound Ip on RSL-150 Content
α- Thujene
α- Pinene
Camphene
Sabinene
Oct-1-en-3-ol
Myrcene
α- Phellandrene
α- Terpinene
p- Cymene
1,8- Cineole
γ- Terpinene
trans- Pinene hydrate (?)
Pinocamphone
Isopinocamphone
Terpinen-4-ol
p- Cymen-8-ol
α-Terpineol
Methyl thymol ether (?)
Cuminaldehyde
Thymol
Carvacrol
α- Terpinyl acetate
Eugenol
α- Ylangene
β- Elemene
Thymohydroquinone
dimethylether
β- Caryophyllene
α- Humulene
β-Farnesene
allo- Aromadendrene
δ- Cadinene
Nerolidol
Spathulenol
Caryophyllene oxide
921
931
941
964
967
981
995
1007
1011
1019
1049
1131
1139
1155
1164
1170
1173
1222
1228
1267
1274
1329
1330
1369
1382
1400
1418
1447
1449
1461
1515
1547
1564
1570
0.1
10.6
0.1
1.5
0.3
0.6
4.1
tr
6.3
6.6
1.1
0.3
1.0
33.5
0.7
0.4
0.6
0.1
tr
0.2
0.3
1.2
tr
tr
tr
16.1
0.9
0.1
0.1
tr
1.2
0.1
0.7
0.4
tr = trace (<0.05%); (?) = identification based on the
MS and RT
Source: Pooter et al. (1991)
Farmers knowledge and perception towards use of S.
suaveolens in insect pests and weeds management
The diversity of insect pests and weeds in most
agricultural lands need a multi-control strategies to
produce satisfactory results in a sustainable manner
(Parker et al., 2013). The goals and values of long-
term sustainability must be reflected in combinations
of practices and methods consistent with an
individual farmer's resources, including knowledge
and farming practices (Ikerd, 1993). Unfortunately,
most smallholder farmers in developing countries
have limited knowledge and are resource-
constrained. This limits their capacity to manage
weeds and insect pests (Whitbread et al., 2010).
Pest management practices by most smallholder
farmers are mainly based on use of chemical
pesticides, though this alone does not give the desired
results (Toda & Morishita, 2009). Few of these
farmers have combined such method with some
cultural practices such as intercropping and crop
rotation (Ajeigbe et al., 2010; Ngowi et al., 2007).
Other studies reported that limited technical
knowledge among small holder famers and shortage
of extension services are among the limiting factors
that hinder the adoption of suitable pest management
practices (Midega et al., 2012; Mkenda et al., 2020).
Most farmers still relying on past experience and
farming practices despite the fact that they have not
attained fruitful results over the years (Khan &
Damalas, 2015). Integrating different pest
management practices has long been proposed as the
long term solution and future for sustainable
agriculture (Pretty & Bharucha, 2015).
For any pest management approach to work and
eventually adopted by farmers, their knowledge,
perceptions and practices has to be fully realized
(Chitere & Omolo, 2008; Hashemi & Damalas, 2010;
Huis, 2014). Khan et al. reported that some farmers are
aware of the role played by companion crops with
repellent or toxic characteristics in pests control, as well
as harboring natural enemies and in this regard, S.
suaveolens may have considerable potential (Khan et al.,
2010). Isman and Grieneisen (2014) pointed out several
plant species from the Asteraceae and other families
with pesticidal properties that may be used to control
and manage insect pests and weeds, however, very few
of these plants are known and used by smallholder
farmers despite the increasingly focus in research on
plant species with pesticidal potential in Africa.
Conclusion
J. Bio. & Env. Sci. 2020
107 | Laizer et al.
S. suaveolens is a weed which possesses diverse group
of biological activities both in medicine as well as in
agriculture.
However, the importance of such activities on the
later have been ignored despite its potential in
managing insect pests in the field, storage and
suppressing other weeds. The wide geographical
distribution of S. suaveolens give an added advantage
and opportunities to small holder farmers as a cheap
alternative in managing weeds and controlling pests
since they cannot afford the synthetic pesticide.
Additionally, the combination of small dosage of
synthetic pesticides with botanical extracts may be more
effective and environmental friendly compared with
standard dose of synthetic pesticides (Isman, 2006b;
Joseph et al., 2008). The allelochemicals from the plant
extracts may be isolated and identified and eventually
serves as templates for developing new generation of
pesticides with less toxic effect to environment and
human health. Extension services and trainings are very
important in enhancing the performance and promoting
adoption of new strategies and practices to smallholder
farmers such as use of botanical extracts particularly
from invasive weeds in managing and controlling pests,
other weeds and diseases.
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Insights of allelopathic, insecticidal and repellent potential of an invasive plant Sphaeranthus suaveolens in pest and weed management | JBES

  • 1. J. Bio. & Env. Sci. 2020 101 | Laizer et al. REVIEW PAPER OPEN ACCESS Insights of allelopathic, insecticidal and repellent potential of an invasive plant Sphaeranthus suaveolens in pest and weed management Hudson Laizer*1,2 , Musa Chacha1 , Patrick Ndakidemi1 1 Department of Sustainable Agriculture and Biodiversity Conservation, Nelson Mandela African Institution of Science and Technology, Arusha, Tanzania 2 Centre for Research, Agricultural Advancement, Teaching Excellence and Sustainability in Food and Nutritional Security (CREATES), The Nelson Mandela African Institution of Science and Technology (NM-AIST), Arusha, Tanzania Article published on August 30, 2020 Key words: Botanical extracts, Secondary metabolites, Sustainable agriculture, Smallholder farmers, Terpene Abstract Sphaeranthus suaveolens is a weed from the family Asteraceae, it grows abundantly in wet areas and most common in rice fields. The extracts from plants closely related to S. suaveolens have been reported to have allelopathic, insecticidal, antifeedant, repellent, and other biological activities. Currently, the use of synthetic chemicals to control weeds and insect pests raise several concerns related to environment and human health. Extracts from plants with pesticidal properties can offer the best and an environmentally friendly alternative. Some of these extracts have been extensively tested to assess their applications as valuable natural resources in sustainable agriculture. This review article therefore explores the potential of S. suaveolens extracts in controlling insect pests and managing weeds by smallholder farmers. *Corresponding Author: Hudson Laizer  laizerh@nm-aist.ac.tz Journal of Biodiversity and Environmental Sciences (JBES) ISSN: 2220-6663 (Print) 2222-3045 (Online) Vol. 17, No. 2, p. 101-112, 2020 http://www.innspub.net
  • 2. J. Bio. & Env. Sci. 2020 102 | Laizer et al. Introduction Agricultural production in most part of the world is affected by pests, diseases and weeds among other factors (Nations, 2016; Singh et al., 2003). Majority of the farmers respond to these constraints through the use of synthetic chemicals such as herbicides and pesticide (Kelly et al., 2003). The extensive use of these chemicals in controlling insect pests and managing weeds, however, have alarmed the public on the effects they might bring to human health and the environment at large (Khanh et al., 2005). Such concerns are putting pressure on agricultural sector to reduce the use of chemicals and as a result, much attention is paid to alternative methods and techniques for controlling and managing weeds and insect pests, through non-chemical methods and/or use of natural products such as botanical extracts (Isman, 2006a; Williamson et al., 2008). Prior to the discovery and commercialization of synthetic pesticides, botanical extracts, among other methods, were used by most farmers in crop protection against insect pests, weeds and diseases (El-Wakeil, 2013). Extracts from plants with allelopathic or pesticidal properties were of great importance in making natural herbicides and pesticides (Benner, 1993; Godfrey, 1994). There is also increasingly evidence from literatures that plant extracts can be manipulated and used as perfect agrochemicals in controlling insect pests and managing weeds (Hoagland, 2001; Macías et al., 2001; Mkenda et al., 2015; Nattudurai et al., 2012; Ngondya et al., 2016; Singh et al., 2003; Stephen et al., 2002; Vyvyan, 2002). The secondary metabolites in plants are responsible to biological activities that offer defense against predators, fungi and bacteria, also these metabolites may act as natural herbicides by suppressing other plant species (Dewick, 2009; Schoonhoven et al., 2005). These biological activities from the plants secondary metabolites can be exploited and manipulated for various human uses, and in this respect Sphaeranthus suaveolens has a considerable potential. S. suaveolens is a widespread weed in swampy and irrigated farmlands, and usually infests cultivated fields and reduces crop productivity (Beentje, 2002). A heavy infestation of this weed results in adverse effects on the growth and yield of crops, particularly in rice fields (Fahmy, 1997). It has been observed that S. suaveolens has an ability to overcome and suppress crop plants in a wide range over a short period of time (Ivens, 1989). However, the secondary metabolites involved are largely unknown and weather they can be applied in managing other weeds and controlling insect pests is yet to be determined. Understanding this could considerably justify the practical application of botanical-based weeds and insect pests management techniques for most smallholder farmers in areas where S. suaveolens is growing. This review article therefore highlights the allelopathic, insecticidal and repellent potential of an invasive plant S. suaveolens with a focus on its application in controlling insect pests and managing weeds by smallholder farmers. Overview of S. suaveolens and its spatial distribution in Tanzania S. suaveolens is an aromatic annual spreading herb from the family Asteraceae with broad sessile leaves covered with glandular hairs (Osman, 2011). The lower stem often trails along the ground and roots at the nodes with thread-like root, flowers are purple, in compound heads ovoid in shape and borne on solitary glandular peduncles with toothed wings (Fayed & Mohamed, 1991). The head as a whole is surrounded by several rows of bracts, of which only the tips are visible when flowers are fully open, and propagated by seeds which takes about 10-12 days to germinate, and the seedlings attain the height of 5.0 - 6.0cm within 30 days in a favorable environment (Beentje, 2002). S. suaveolens is widespread in Africa over a range of altitudes from Rwanda, Burundi, Sudan, Ethiopia,
  • 3. J. Bio. & Env. Sci. 2020 103 | Laizer et al. Zambia, Malawi, Mozambique, Egypt, Uganda, Kenya to Tanzania (Everard et al., 2002; Fahmy, 1997). It is mostly found growing in wet areas and thrives well in medium clayey soils, but also common in and around irrigation ditches and rice fields and considered as a major weed in most farms (Ivens, 1989). In Tanzania, the plant was first reported in Songea (Brenan, 1960), then Mpwapwa (Launert, 2003), Mkata and Mandela in Wami River Ecosystem (Mligo, 2017). The spatial distribution of S. suaveolens in Northern Tanzania particularly Arusha and Kilimanjaro regions however, have not been mapped despite being reported as a weed to most agricultural fields. Fig. 1. Distribution of S. suaveolens in Wami River Ecosystem in Tanzania. Allelopathic effects of S. suaveolens on crops Allelopathy is a phenomenon, whereby one plant influences the growth of another one, including microorganisms by the release of chemical compounds into the environment (Keeley, 2010; Rice, 1983; Whittaker & Feeny, 1971). The allelopathic effects are the result of chemical compounds known as allelochemicals, which are usually plants’ secondary metabolites or byproducts of the principal metabolic pathways in plants (Chancellor, 1987; Dayan et al., 2009; Macías et al., 2007). In recent years, allelopathy has become a research hotspot for making comprehensive analysis about the mechanism of weeds and identification of specific chemical compounds responsible for allelopathic effects (Azirak & Karaman, 2008; Bais et al., 2003; Einhellig & Leather, 1988). Most allelopathic plants have been observed to significantly affect the growth, productivity and yield of other crops by causing soil sickness and nutrient imbalance (Kohli et al., 2008), as well as affecting the microbial population (Batish et al., 2001). Several studies have indicated that, most weeds possess allelopathic effects which play a significant roles in their invasion success (Macías et al., 2014; Qasem & Foy, 2001; Zhou et al., 2013). Numerous weeds from the Asteraceae family have been reported to possess allelopathy and can significantly inhibit crop productivity in agricultural land (Ilori et al., 2010; Kong et al., 2007). The invasive weed, Sphaeranthus indicus has been reported to inhibit seed germination and growth of wheat (Triticum aestivum), rice (Oryza
  • 4. J. Bio. & Env. Sci. 2020 104 | Laizer et al. sativa) and mung bean (Vigna radiata) in different farming systems (Lodha, 2004). Recently, Mahajan et al. (2015) reviewed the allelopathic potential of S. indicus and found that the germination and seedling growth of various crops were significantly decreased with increase in concentration of its extract. Despite of the negative effects on cultivated crops, allelochemicals from allelopathic plants can be manipulated and used to control weeds of various crops. For example, Khanh et al. (2006) noted that the allelochemicals contained in tissues of Passiflora edulis can significantly suppress the two noxious paddy rice weeds (Echinochloa crusgalli and Monochoria vaginalis). Other studies have also reported the use of allelochemicals for weed control in the laboratory as well as application under field conditions (Jabran et al., 2015; Ngondya et al., 2016). Jamil et al. (2009) described the utilization of allelopathic water extract as an important and useful way of exploiting the allelopathic potential to manage wild oat and canary grass in wheat fields. The emergence and root length of most rice weeds was inhibited by allelochemical (Lycorine) from the dead leaves of spider lily (Iqbal et al., 2006). The allelopathic crops when used as cover crop, mulch, smother crops, green manures, or grown in rotational sequences maybe helpful in reducing noxious weeds and plant pathogen, improve soil quality and crop yield (Khanh et al., 2005). Furthermore, the application of allelopathic extracts may give an efficient alternative control over weeds similar to that offered by synthetic herbicides (Xuan et al., 2004). Interactions among potential allelopathic plants, target pests and other non- target organisms in a cropping system also need to be considered and fully realized to avoid detrimental effects to desired crops and non-target species (Farooq et al., 2013). The allelochemicals involved in weed suppression can serve as basic templates for developing new generation of biopesticides with low or no toxic effects to the environment and human health (Ferguson et al., 2009). The allelopathic potential of S. suaveolens, therefore need to be realized and selectively used to suppress density of other weeds and insect pests population particularly in small farming agricultural systems. Insecticidal and repellent activities of S. suaveolens to insect pests Insecticides, weather natural or synthetic are developed to either kill, repel, or interfere with the damaging behavior of insect pests (EPA, 2009). Due to intensity of plant-insect interactions, plants have well developed defense mechanisms against insect pests by producing natural compounds which acts as natural pesticides (Després et al., 2007). The most exciting concept is to isolate and identify such compounds and use them as candidates in making safer pesticides (Maia & Moore, 2011). Plant extracts with pesticidal properties can be active against specific target insects, biodegradable, have low to non-toxic effects, cheap and easy to prepare (Kim et al., 2003; Mkindi et al., 2017). Due to this facts, these plant extracts could lead to the development of new classes of safer pesticides (Céspedes et al., 2014; Tembo et al., 2018). The Asteraceae family in which S. suaveolens belongs, have been reported to contain plants with insecticidal activities (Dhale, 2013; Green et al., 2017; Sosa et al., 2018). These insecticidal activities are mostly linked to presence of secondary metabolites such as terpenes, which can act as larvicides, insect growth regulators and feeding and oviposition deterrents (Miresmailli & Isman, 2014). Terpene is among the most diverse class of plant secondary metabolites found in essential oils of most plants, S. suaveolens included (Ahmed et al., 2017; Pagare et al., 2015). This secondary metabolite has been reported to play an important role in plant protection against pathogens (Neerman, 2003), insects (Wu et al., 2016) and toxic to mammals as well (Gurib-Fakim, 2006). Recently, Sosa et al. (2018) reported insecticidal activities of the terpene isolated from Vernonanthura nebularum against fall army warm (Spodoptera
  • 5. J. Bio. & Env. Sci. 2020 105 | Laizer et al. frugiperda) and fruit fly (Ceratitis capitate). Moreover, two other terpenes from Inula helenium were examined by Kaur et al. (2017) and reported to significantly inhibit the growth of tobacco leafworm (Spodoptera litura). Other techniques such as use of water extracts from S. indicus was also reported to demonstrated toxic effects against insect pests such as rice weevil (Patole et al., 2008), cowpea weevil (Singh & Shrivastava, 2012) and red flour beetle (Pugazhvendan et al., 2012). Furthermore, the extracts from S. indicus have been showing larvicidal activities and repellent activities to most of the insect pests (Arivoli et al., 2016; Baby, 1994; Singh & Shrivastava, 2012). The presence of secondary metabolites with pesticidal properties such as terpene in the essential oil of S. suaveolens may give positive insecticidal and repellent activities to most insect pests, hence used for as protective agents against insect pests, but this needs further scientific investigation. Key allelochemicals from the leaf, stem and root extracts of S. suaveolens Allelochemicals are secondary metabolites produced by living organisms such as plants that have stimulatory or inhibitory effects upon the growth, health, behavior and distribution of neighboring organisms being another plants, insects or microbes (Haig, 2008). The role played by secondary metabolites is mostly ecological, linked to plant defense against other plants, pests, or diseases (Ramakrishna & Ravishankar, 2011). Allelochemicals undoubtedly pose problems in agriculture, but if well manipulated they can be beneficial and offer great opportunities such as insect pests and weeds control (Einhellig, 1987). Despite the efforts in allelopathic researches, little is known on the potential to exploit the key allelochemicals in agricultural systems and use them as templates in making safer and affordable herbicides and/or pesticides (Kremer & Ben- Hammouda, 2009). Much of the work to date has focused on weather extracts from S. suaveolens show biological activities such as antimicrobial, immune stimulating, anticancer, antitumor, anthelmintic, repellency, insecticidal and allelopathy (Ahmed & Mahmoud, 1997; Kleinowski et al., 2016). However, very few literatures have reported the identified compounds found in S. suaveolens extracts, none of it has a list of allelochemicals found in S. suaveolens despite being the weed of economic importance in many rice and common bean farms in Africa. Allelochemicals belong to various chemical groups, and can be classified based on their structures and properties into: water-soluble organic acids, straight- chain alcohols, aliphatic aldehydes, and ketones, lactones, long-chain fatty acids and polyacetylenes, quinines (benzoquinone, anthraquinone and complex quinines), phenolics, cinnamic acid and its derivatives, coumarins, flavonoids, tannins, steroids and terpenoids (Li et al., 2010). Most of these biochemicals are synthesized during the shikimate pathway (Hussain & Reigosa, 2011) or, in the case of essential oils, from the isoprenoid pathway (Rehman et al., 2016). The extract of the aerial parts of S. suavealens was reported by Jakupovic et al. (1990) to contain eight eudesman-12. 6 β abides, carvotacetone derivatives and a thymohydroquinone glucopyranoside. Later on, Pooter et al. (1991) reported extract of the same plant comprise of thymohydroquinone dimethylether, a diacetylene thiophene, inositol and myoinositol esters, and several carvotanacetone derivatives, he went further to examine the essential oil of S. suaveolens and noted methyl chavicol, α-ionon,e, dcadinene and p- methoxycinnamadehyde as major constituents, and a- terpinene, citral, geraniol, geranyl acetate, βionone, shaerene, indicusene and sphaeranthol as minor constituents. Ahmed and Mahmoud (1997) examine the extract of aerial parts of S. suaveolens and reported three carvotacetone derivatives, together with four monoteroene compounds. Later on, Hassanali et al. (1998) reported cis-pinocamphone as the major constituents (63.5%) of the leaf oil of S. sauveolens. The details of these identified compounds are stated
  • 6. J. Bio. & Env. Sci. 2020 106 | Laizer et al. in Table 1, however, identifying alone isn’t sufficient enough, rather gaining an understanding on which among these compounds are allelochemicals and how to use them in improving crop production though managing weeds and controlling insect pests in sustainable agriculture will be a big advantage. Table 1. Identified compounds from leaf oil of S. suaveolens. Compound Ip on RSL-150 Content α- Thujene α- Pinene Camphene Sabinene Oct-1-en-3-ol Myrcene α- Phellandrene α- Terpinene p- Cymene 1,8- Cineole γ- Terpinene trans- Pinene hydrate (?) Pinocamphone Isopinocamphone Terpinen-4-ol p- Cymen-8-ol α-Terpineol Methyl thymol ether (?) Cuminaldehyde Thymol Carvacrol α- Terpinyl acetate Eugenol α- Ylangene β- Elemene Thymohydroquinone dimethylether β- Caryophyllene α- Humulene β-Farnesene allo- Aromadendrene δ- Cadinene Nerolidol Spathulenol Caryophyllene oxide 921 931 941 964 967 981 995 1007 1011 1019 1049 1131 1139 1155 1164 1170 1173 1222 1228 1267 1274 1329 1330 1369 1382 1400 1418 1447 1449 1461 1515 1547 1564 1570 0.1 10.6 0.1 1.5 0.3 0.6 4.1 tr 6.3 6.6 1.1 0.3 1.0 33.5 0.7 0.4 0.6 0.1 tr 0.2 0.3 1.2 tr tr tr 16.1 0.9 0.1 0.1 tr 1.2 0.1 0.7 0.4 tr = trace (<0.05%); (?) = identification based on the MS and RT Source: Pooter et al. (1991) Farmers knowledge and perception towards use of S. suaveolens in insect pests and weeds management The diversity of insect pests and weeds in most agricultural lands need a multi-control strategies to produce satisfactory results in a sustainable manner (Parker et al., 2013). The goals and values of long- term sustainability must be reflected in combinations of practices and methods consistent with an individual farmer's resources, including knowledge and farming practices (Ikerd, 1993). Unfortunately, most smallholder farmers in developing countries have limited knowledge and are resource- constrained. This limits their capacity to manage weeds and insect pests (Whitbread et al., 2010). Pest management practices by most smallholder farmers are mainly based on use of chemical pesticides, though this alone does not give the desired results (Toda & Morishita, 2009). Few of these farmers have combined such method with some cultural practices such as intercropping and crop rotation (Ajeigbe et al., 2010; Ngowi et al., 2007). Other studies reported that limited technical knowledge among small holder famers and shortage of extension services are among the limiting factors that hinder the adoption of suitable pest management practices (Midega et al., 2012; Mkenda et al., 2020). Most farmers still relying on past experience and farming practices despite the fact that they have not attained fruitful results over the years (Khan & Damalas, 2015). Integrating different pest management practices has long been proposed as the long term solution and future for sustainable agriculture (Pretty & Bharucha, 2015). For any pest management approach to work and eventually adopted by farmers, their knowledge, perceptions and practices has to be fully realized (Chitere & Omolo, 2008; Hashemi & Damalas, 2010; Huis, 2014). Khan et al. reported that some farmers are aware of the role played by companion crops with repellent or toxic characteristics in pests control, as well as harboring natural enemies and in this regard, S. suaveolens may have considerable potential (Khan et al., 2010). Isman and Grieneisen (2014) pointed out several plant species from the Asteraceae and other families with pesticidal properties that may be used to control and manage insect pests and weeds, however, very few of these plants are known and used by smallholder farmers despite the increasingly focus in research on plant species with pesticidal potential in Africa. Conclusion
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