Toxicon 193 (2021) 21–27

Contents lists available at ScienceDirect

Toxicon
journal homepage: http://www.elsevier.com/locate/toxicon

Review

Saponin toxicity as key player in plant defense against pathogens

Madiha Zaynab a, b, c, Yasir Sharif d, Safdar Abbas e, Muhammad Zohaib Afzal f,
Muhammad Qasim g, Ahlam Khalofah h, i, Mohammad Javed Ansari j, Khalid Ali Khan h, i, k,
Li Tao l, Shuangfei Li b, *
a
College of Physics and Optoelectronic Engineering, Shenzhen University, Shenzhen, 518060, China
b
Shenzhen Key Laboratory of Marine Bioresource & Eco-environmental Sciences, College of Life Sciences and Oceanography, Shenzhen University, Shenzhen,
Guangdong, 518071, China
c
Shenzhen Environmental Monitoring Center, Shenzhen, 518049, Guangdong, China
d
College of Plant Protection, Fujian Agriculture and Forestry University, Fuzhou, 350002, Fujian, China
e
Department of Biochemistry, Faculty of Biological Sciences, Quaid-i-Azam University, Islamabad, Pakistan
f
Department of Plant Breeding and Genetics, University of Agriculture, Faisalabad, 38000, Pakistan
g
Ministry of Agriculture Key Lab of Molecular Biology of Crop Pathogens and Insects,Institute of Insect Science,Zhejiang University, Hangzhou, 310058, China
h
Department of Biology, Faculty of Science, King Khalid University, P.O. Box 9004, Abha, 61413, Saudi Arabia
i
Research Center for Advanced Materials Science (RCAMS), King Khalid University, P.O. Box 9004, Abha, 61413, Saudi Arabia
j
Department of Botany, Hindu College Moradabad (MJP Rohilkhand University Bareilly), 244001, India
k
Unit of Bee Research and Honey Production, Faculty of Science, King Khalid University, P.O. Box 9004, Abha, 61413, Saudi Arabia
l
Shenzhen Base of South China Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Shenzhen, China

A R T I C L E I N F O A B S T R A C T

Handling Editor: Kevin Welch Microbial pathogens attack every plant tissue, including leaves, roots, shoots, and flowers during all growth
stages. Thus, they cause several diseases resulting in a plant’s failure or loss of the whole crop in severe cases. To
Keywords: combat the pathogens attack, plants produce some biologically active toxic compounds known as saponins. The
Bacteria saponins are secondary metabolic compounds produced in healthy plants with potential anti-pathogenic activity
Insects
and serve as potential chemical barriers against pathogens. Saponins are classified into two major groups the
Secondary metabolites
steroidal and terpenoid saponins. Here, we reported the significance of saponin toxins in the war against insect
Natural products
Triterpenoid pests, fungal, and bacterial pathogens. Saponins are present in both cultivated (chilies, spinach, soybean, quinoa,
onion, oat, tea, etc.) and wild plant species. As they are natural toxic constituents of plant defense, breeders and
plant researchers aiming to boost plant imm
unity should focus on transferring these compounds in cash crops.

1. Introduction secondary metabolites have major roles in reacting to particular stress

Almost one-third of the 450,000 discovered plant species of this
world are at the risk of extinction (Pimm and Joppa, 2015). The
approximate cumulative amount of plant metabolites in a species ex­
ceeds 10,000, but presently only fewer than 20% of these metabolites
can be analyzed using metabolome methods (Abdelrahman et al., 2018;
Lei et al., 2011). Throughout human history, people have often been
using natural plant extracts for centuries as traditional medicines.
Generally speaking, plants synthesize various primary and secondary
metabolites with varying structures (Arbona et al., 2013). The primary
metabolites are essential for plant development and growth, and
for plants (Abdelrahman et al., 2019; Fujii et al., 2015). The secondary
metabolites’ defensive functions have been confirmed by an in vitro
examination of plants (Mes et al., 2000). The production of secondary
metabolites results from millions of years of interaction between plants
and pathogens and is thought to include more than 100,000 metabolites,
but their interaction needs more clarification (Wink, 2008). Although
plants with a high concentration of secondary metabolites are regarded
as more resistant to biotic and abiotic stresses, their development has
been deemed to be costly for plant growth and reproduction (Karban
et al., 1997; Simms, 1992). The structure and function of secondary
metabolites indicate why the induced defense has evolved from plants,

* Corresponding author.
E-mail address: szu_sfli@163.com (S. Li).

https://doi.org/10.1016/j.toxicon.2021.01.009
Received 4 November 2020; Received in revised form 24 December 2020; Accepted 20 January 2021
Available online 26 January 2021
0041-0101/© 2021 Elsevier Ltd. All rights reserved.
M. Zaynab et al.
which is typified by a rise in stress levels (Tollrian and Harvell, 1999).
Several studies have shown that hundreds of plant compounds have
protective, ecological, and chemical functions (Harborne, 1989).
Saponin compounds have a remarkable chemical composition and bio­
logical function. Saponins production is part of a plant’s natural growth
and developmental process, while they also act as significant chemical
obstacles to pathogenic fungal and insect resistance mechanisms
(Abdelrahman et al., 2017). Plants are highly capable of producing
many secondary metabolites, including alkaloids, glycosides, terpe­
noids, saponins, and steroids (Khan et al., 2018; Mujeeb et al., 2014;
Othman et al., 2019). Many natural products are incredibly successful
fighters against pathogens (Cragg and Newman, 2013; Rossiter et al.,
2017; Salam and Quave, 2018). Saponins are a heterogeneous group of
secondary metabolites, distinguished by the presence of a structure with
a steroid or triterpenoid aglycone and one or more sugar chains (Moses
et al., 2014). They are inactive precursors in plant cells but readily
transformed to biologically active antibiotics by plant enzymes’ activity
(Güçlü-Üstündağ and Mazza, 2007; Hostettmann and Marston, 1995;
Osbourn, 1996). Such compounds may also be interpreted as "pre­
formed" as they are already present in stable plant tissues with plant
enzymes that activate them (Osbourn, 1996). They are also utilized in
drugs, flavor modification, and cosmetics (Hostettmann and Marston,
1995; Kregiel et al., 2017). These are typically considered to play a
significant role in the defense due to their antimicrobial, antifungal,
insecticidal, and anti-feeding properties against bacteria and herbicides
(Augustin et al., 2011; Kregiel et al., 2017; Morrissey and Osbourn,
1999; Osbourn et al., 2011; Sparg et al., 2004; Trdá et al., 2019).
Saponin levels enhance response to stresses, regulated via a dynamic
signaling cascade involving jasmonate and salicylate hormones through
transcriptional stimulation of biosynthetic genes (de Costa et al., 2013).
Therefore, elicitors stimulate the biosynthesis of such molecules. This
trait has been used in many plant species to increase the saponin yield
(Shabani et al., 2009; Yendo et al., 2010).
2. Types and classification of saponins
The saponins are a family of substances with a compact structure
composed of four hydrocarbon rings to which sugar is bound in groups
of one or two (generally not more than ten units). Saponins comprise a
large class of plant-based natural products, including steroids (C27),
triterpene (C30), and glycosides, which are further divided into 11 main
carbon-skeleton-based subgroups (Bailly and Vergoten, 2020). These 11
subgroups are cucurbitanes, lanostanes, cycloartanes, ursanes, tarax­
asteranes, oleananes, hopanes, lupanes, steroids, tirucallanes, and
dammaranes (Kregiel et al., 2017). Many forms of saponins arise from a
combination of glycones and carbohydrates in different attachment
positions. The aglycone’s molecular characteristics (known as sapo­
genin) separate the saponins into steroidal and triterpenoid saponins
(Qasim et al., 2020). Steroidal saponin is primarily a compound of 27
carbon atoms with a central structure: spirostan and furostan (Sparg
et al., 2004). The terpenoids’ saponins mainly comprise aglycones with
30C atoms or their non-derivatives (Vincken et al., 2007). Epoxy groups,
keto functions, and double bonds between C-12 and C-13 have also been
reported. The classes of hydroxyl may be acylated, contributing to the
development of ester saponins. Acid products are most commonly acetic,
formic, a-methyl butyric, n- and iso-butyric, angelic, ferulic acid, cin­
namic, benzoic, tiglic, and a methyl-butyric in some instances, sulphuric
acid, tiglic, benzoic, cinnamic, ferulic acid (Hill and Connolly, 2011;
Zhao and Shi, 2011; Zhou et al., 2011). The saponins carbohydrate
chains usually include D-glucose, D-glucuronic acid, D-fucose, D-apiose,
D-galactose, L-arabinose, D-xylose, and L-rhamnose. The sugars often
bind C-3 as glycosides or carboxylic esters in position C-28 through the
hydroxyl ring (Arslan, 2014; Thakur et al., 2011). The saponin’s
chemical structure may also be transformed in the processing or storage
time. The relationship between the sugar chain and the aglycones and
between the sugar residues may be hydrolyzed by hydrothermolysis or
22
Toxicon 193 (2021) 21–27
enzyme/microbial transformations, resulting in aglycones’ production
prosapogenins (in part hydrolyzed saponins) and sugar residues (Oles­
zek and Hamed, 2010; Hossain, 2016).
A significant number of plant families synthesize saponin (Dinda
et al., 2010; Hussain et al., 2019; Man et al., 2010; Sparg et al., 2004;
Vincken et al., 2007). Cultivated and wild plants such as Quillaja, sugar
beet, oats, legumes, ginseng, asparagus, and alfalfa have saponin com­
pounds (Badenes-Perez et al., 2014). Saponins are generally categorized
into three major classes in the plant kingdom; steroidal, triterpenoid,
and steroidal glycoalkaloid (Vincken et al., 2007). Triterpenoid saponins
are an essential part of the plant defense system and are primarily found
in various crops. Several prominent plant families include Zygophylla­
ceae, Myrsinaceae, Chenopodiaceae, Berberidaceae, Cucurbitaceae, Aral­
iaceae, Aquifoliaceae, Caryophyllaceae, Apiaceae, Amaranthaceae, and
Leguminosae (Hostettmann and Marston, 1995; Kregiel et al., 2017;
Mert-Türk, 2006; Parente and da Silva, 2009; Shi et al., 2004; Sparg
et al., 2004) have triterpenoid saponins. Similarly, various plant sections
have specific saponin concentrations and types (Zhou et al., 2018). For
instance, in the same field, a leaf produces more types and amounts of
saponins than shoots or flowers. The age and components of plants
determined the kind of saponin molecules (Wei et al., 2018). Di­
cotyledons plants, such as Quillaja bark (Fleck et al., 2019), Primula roots
(Katselis et al., 2007; Koczurkiewicz et al., 2015; Park et al., 2005;
Schmid et al., 2018; Sundaramoorthy et al., 2019; Yoshikawa et al.,
1996), Ginseng roots, Polygala Amarae leaves, glycyrrbizae roots, Senega
roots, Glycine max seeds, Herniaria leaves, and Saponariae roots are rich
sources of triterpenoids (Coran and Mulas, 2012; De Geyter et al., 2007a;
Estrada et al., 2000; Kimura et al., 2006; Kregiel et al., 2017; Lu et al.,
2015; Sundaramoorthy et al., 2019).
However, the quality of saponin molecules due to fluctuation of
environmental conditions differs significantly in quantity and type
(Phrompittayarat et al., 2011). Different cultivated plants are fortified
with numerous saponins that are responsible for plant safety (Singh
et al., 2017). Solanaceous plants (e.g., potatoes and tomatoes) are often
well-enriched by numerous steroidal glycoalkaloids that are important
for herbivores and microbes resistance (Moses et al., 2014). On the other
hand, many steroid glycosides are also a popular component of many
crops, beneficial for the plant immune system, such as Trigonella foe­
num-graecum, S. Lycopersicum, Solanum melongena, Panax ginseng, Dio­
scorea spp, Capsicum spp, Avena sativa, Asparagus Officinalis, and Allium
spp. The ultimate industrial source of triterpenoid and steroidal sapo­
nins are commonly recognized as Yucca plant (Singh et al., 2017). Ce­
reals and grasses are generally deficient in saponins, with some notable
exceptions, such as the Avena species (oats), accumulating both tri­
terpenoid and steroidal saponins (Osbourn, 2003; Price et al., 1987;
Yang et al., 2016).
The steroid saponins are usually found almost exclusively in the
Scrophulariaceae, Palmae, Bromeliaceae, Agavaceae, Amaryllidaceae, Lil­
iaceae, Dioscoreaceae, Asparagaceae, and Alliaceae families but also in the
dicotyledonous Solanaceae family (Table 1.) (Kregiel et al., 2017; Moses
et al., 2014). In crop plants, steroid saponins are accumulated in abun­
dance in ginseng, yucca, fenugreek, asparagus, alliums, and yams.
Functionally, some steroidal saponins are lethal to many harmful soil
microbes as they are present in plant roots abundantly (Faizal and
Geelen, 2013). Steroidal saponins have also been reported in the leaves
of Dioscorea caucasica (Gurielidze et al., 2004). This review article is
based on the plant-related saponins’ role against plant pathogens.
3. Toxicity against pathogens
The biosynthesis of secondary metabolites in response to biotic or
abiotic stresses is a challenge. Production of saponin toxin is a cost-
effective strategy for combatting pathogens, regardless of external in­
dications, leading to innate immunity. The drawback of saponin ag­
gregation is that they absorb a large amount of energy and that
pathogens may improve their immunity. It is avoided as saponin
M. Zaynab et al. Toxicon 193 (2021) 21–27

Table 1 bacterial strains; for Fr1, the MIC values were lower than Fr2 and Fr3.
Types of saponins present in different plant families. Owing to specific forms of main saponins, the inhibition zone (I.Z.)
Species Saponin type Family References values of Fr1 were significantly higher. Likewise, the Fr1 demonstrated
fairly higher bacterial lethality and viability (Khan et al., 2018). Sapo­
Celosia cristata Triterpenoid Amaranthaceae Wang et al. (2010)
Aralia elata Triterpenoid Araliaceae Lee et al. (2009a) nins resist strongly against S. Aureus (gram-positive bacteria) and
Physospermum Triterpenoid Apiaceae Tundis et al. (2009) moderately against gram-negative bacteria (E. coli and S. enteritidis). Fr1,
verticillatum therefore, displayed comparatively low in vitro antibacterial activity. Fr1
Ilex kudingcha Triterpenoid Aquifoliaceae Zuo et al. (2012) saponins were considered successful in the dose-dependent analysis of
Silphium radula Triterpenoid Asteraceae Calabria et al. (2008)
Tacca integrifolia Steroidal Taccaceae Shwe et al. (2010)
released soluble proteins and APK contents in destroying bacterial cell
Smilax excelsa Steroidal Smilacaceae Ivanova et al. (2009) wall. Saponin damaged the wall of bacteria with various potential for
Anemarrhena Steroidal Liliaceae Lee et al. (2010) each strain of bacteria. Lysozyme is a component of normal and
asphodeloides non-specific immunity. The actions of molecules guarding against bac­
Panicum virgatum Steroidal Poaceae Lee et al. (2009b)
terial attacks are considered an effective innate defense agent in plants
Dioscorea nipponica Steroidal Dioscoreacea Dioscorea nipponica
Asparagus fifilicinus Steroidal Asparagaceae Wu et al. (2010) (Riera Romo et al., 2016). Tagousop et al. (2018) reported the antimi­
Agave utahensis Steroidal Agavaceae Yokosuka and Mimaki crobial activities of Melanthera elliptica saponins and their synergistic
(2009) action with antibiotics against pathogens’ phenotypes. An unprece­
Cestrum diurnum Steroidal Solanaceae Fouad et al. (2008) dented saponin is being reported herein together with five known
Achras sapota Triterpenoid Sapotaceae Ahmed et al. (2008)
Fadogia ancylantha Triterpenoid Rubiaceae Mencherini et al.
compounds from the methanol extract of the roots of Albizia adianthi­
(2010) folia. The metabolites were obtained using repeated open column and
Actaea racemosa Triterpenoid Ranunculaceae Cicek et al. (2010) spectroscopic chromatography methods followed by spectrometric
Androsace umbellata Triterpenoid Primulaceae Wang et al. (2008) techniques. Eleven gram-negative bacteria were screened for isolated
Polygala crotalarioides Triterpenoid Polygalaceae Hua et al. (2010)
compounds. Saponins extracted from Melanthera elliptica have antibac­
terial activity. The effects of Quillaja saponaria saponins on prokaryotic
precursors stack up, and the saponin content decreases due to chemical and eukaryotic cells have been analyzed by Sonfack et al. (2019). The
changes of the precursor molecules stimulated. In response to pathogen antibacterial effect of ampicillin, streptomycin, and ciprofloxacin was
attack (Morrissey and Osbourn, 1999), saponin contents may increase measured by cultivation methods in the presence of saponins. Saponins
(Szakiel et al., 2011) due to partial or full hydrolyses of stored precursors extracted from Q. saponaria showed cytotoxic effects (Arabski et al.,
in plants or regulated pathogen degradation. Oat has been a model in the 2012). Crude saponin extracts from the aerial component of Anabasis
study of saponin biosynthesis as part of bacteria and fungi interaction articulata were studied utilizing the disc diffusion methods for antibac­
(Mugford et al., 2009; Osbourn, 1996, 2003). The two forms of saponins, terial infection. Five bacterial species (gram-negative bacteria and
steroidal avenacosides, and triterpene avenacins diecoslides, are present gram-positive) were studied. Saponins extract was active with the con­
in the leaves and roots. Avenacins are the components of the active centrations ranged from 0.5 to 1 mg/ml, against all bacteria checked
glycolytic pathway, accumulating in root tip epidermal cells, have (Pseudomonas aeruginosa ATCC 14028, Klebsiella pneumonia, 6633, Ba­
strong antifungal behavior, which offers a wide range of resistance cillus subtilis ATCC, Staphylococcus aureus ATCC 6538, Escherichia coli
against soil born pathogens (Osbourn, 1996). Biologically active con­ ATCC 25922). Phytochemical studies revealed that plants are especially
centrations are often emitted into the soil rhizosphere by an unexplained abundant in saponins that could be induced by anticandidal action
process (Haralampidis et al., 2002; Papadopoulou et al., 1999). The (Maatalah et al., 2012). The antimicrobial efficacy of Gymnema Sylvestre
vacuole contains an active avenacin A-1, and a fluorescence microscope and Eclipta Prostrata leaves saponin fractions under in vitro conditions
is used to detect significant UV fluorescent compounds. The improved has been tested against pathogenic bacteria.
vulnerability of the disease in avenacin-free oat variants confirms sa­
ponin’s significance in pathogen tolerance as phytoprotectants. More­ 3.2. Toxic effect of saponin against fungi
over, intermediate avenacin-biosynthesis pathways accumulation in oat
roots resulted in the callose accumulation, a well-known defense Saponins are known primarily to be part of the antimicrobial pro­
mechanism, which indicates that intermediate accumulation also trig­ tection mechanism of plants. The source of the processes underlying
gers other defense mechanisms (Mylona et al., 2008). their behavior is their capacity to produce complexes with sterols in the
microorganism’s membrane and induce disruption of the membrane
(Augustin et al., 2011; Morrissey and Osbourn, 1999; Sreij et al., 2019;
3.1. Toxic effect of saponin against bacteria Steel and Drysdale, 1988). Saponins’ primary mode of action against
fungi involves the formation of pores and membrane integrity loss.
Natural products are relatively safe, efficient, and essential materials Antifungal activity of saponins for plants has been recognized and
for multipurpose applications and usage. Oleszek et al. (2000) reported documented for decades (Gruiz, 1996; Wolters, 1966). The action
the antibacterial activity of saponins from some plant sources (Hos­ mechanism is similar to saponins’ hemolytic activity. A pattern of action
tettmann and Marston, 1995; Oleszek, 2000). The saponins’ antimi­ was suggested for the avenacin triterpene saponin of oat. The first step
crobial activity extracted from oats, Solanum, Capsicum annuum seeds, involves inserting aglycones in the membrane and then binding them to
garlic, Quillaja, Yucca, and alfalfa have been studied. Saponin extracts sterols (Korchowiec et al., 2015). The following stage contributes to the
were checked against various microbes, including gram-negative and association of sugar residues and sterol-saponin complex formation. It
gram-positive bacteria. Saponins have a detergent potential and can also contributes to membrane lipid re-arrangement, pores formation,
enhance the bacterial cell membrane permeability, which can contribute and, finally, cell lysis (Coleman et al., 2010; Lacaille-Dubois and Wag­
to antibiotic infiltration by the bacterial cell membrane (Jacob et al., ner, 1996). Minutoside saponins and sapogenins, allliogenine, and
1991). Khan et al. (2018) studied green tea’s antibacterial actions neoagigenine extracted from the Allium minutiflorum bulbs are antimi­
(Camellia sinensis) against gram-negative and gram-positive bacteria. crobial against multiple pathogens (Barile et al., 2007). The extraction of
Kim et al. (2015) reported the antimicrobial activities for crude saponins saponin Alliospiroside from Allium cepa is an anti-colletotrichum
extracted from C. sinensis seeds. Khan et al. (2018) isolated saponins gloeosporioids protected strawberry plant, indicating the potential to
from green tea in different fractions by HPLC. They found that Fr1 control plant anthracnose (Teshima et al., 2013). Aescin’s dual mode of
(fraction 1) showed the highest antibacterial activity than Fr2 and Fr3. action incorporating antimicrobial and inductive immune responses
Green tea saponins had different antibacterial effects against various culminated in a very effective blackleg disease inhibition at B. Napus.

23
M. Zaynab et al. Toxicon 193 (2021) 21–27

Aescin, as a potent inducer, induce tolerance in plants similar to BTH or They also carry out healthy insecticide activity by developing choles­
metconazole. Avenacins have a robust antifungal effect (Turner, 1953) terol complexes that cause cell toxicity and insect ecdysial failure
as several other saponins, and triterpenoid avenacin saponins have a (Taylor et al., 2004). Insects can not synthesize sterole structures, so
significant role in the oats against root-infecting fungus Gaeumanno­ cholesterol or phytosterols are essential precursors to ecdysteroid
myces graminis. G. graminis also causing the disease known as a "take-all biosynthesis from food. Saponins produce an indigestible food attach­
disease." and source of considerable yield loss in wheat and barley. The ment of sterols, sterol absorption obstruction, and insect molting and
resistance of oats to G. graminis is due to the presence of saponins in oat ecdysis disturbed. Luciamin insecticide efficacy against the wheat aphid
roots. The central avenacin A-1 is found in an epidermal cell layer of the (Schizaphis graminum) were tested. The toxicity tested by incorporating
oat root tips and even the lateral root initials, thus rendering it suitable luciamin in artificial diets resulted in a constant decrease in aphids’
for providing a chemical barrier to intruding soil-borne microbes, survival. The study’s findings showed a clear deterrent impact of
causing harm to plant tissue (Osbourn et al., 1994; Turner, 1961). Tritici luciamins (500 mM) in wheat aphids and were the first spirostanic sa­
is the existence of such saponins in oat roots (Turner, 1953). Avenacin’s ponins recorded to have insecticidal efficacy (De Geyter et al., 2007b).
role in disease resistance has been shown by investigating oat varieties Nielsen et al. (2010) reported a saponin playing a role in Barbarea vul­
that differ in saponin content. The naturally occurring avenacin content garis resistance against flea beetle (Phyllotreta nemorumi L.). They find
of the Avena species ranges very little (Osbourn et al., 1994; Turk et al., that hederagenin cellobioside is a defensive compound after comparing
2005). However, it has been shown that at least one diploid oat (Avena oleanolic acid cellobioside and hederagenin cellobioside. Studies have
longiglumis) lacks avenacin A-1 and is significantly more sensitive to shown that the insecticidal behavior of saponins depend on the aglycone
fungal conditions than its parents who produce avenacin. It is believed structure and saccharide chain presence. The carbohydrate moiety
that α-tomatine is present in tomato leaves, and this can be shield the removal from aglycone (hederagenin and oleanolic acid) has contrib­
tomato leaves from fungal pathogens. uted to bioactivity losses. Weevils are the primarily stored food pests in
food grains that have been preserved and are usually present in seeds,
nuts, beans, and grain items in dry foods. Several reports have docu­
3.3. Saponin toxicity against insects
mented the usage of saponins to protect the stored grains against wee­
vils. The insecticidal activity of Fenugreek seeds for bean weevil
Plants have also evolved various ways of utilizing secondary me­
evaluated by Pemonge et al. (1997). Triterpenoid saponins isolated from
tabolites to fight against insects (War et al., 2012). An important source
Catunaregam Spinosa have deterrent activity against diamondback moth
for biopesticides development, healthy and sustainable agriculture may
(Gao et al., 2011). All the saponins examined were deterrent or repellent
be searching for plant-derived compounds (Silva et al., 2012). Several
in nature. In tropical countries, the Spodoptera littoralis infestation is
plant secondary metabolites are recognized as feeding deterrents to
popular in vegetable crops (Kregiel et al., 2017; Maia and Moore, 2011).
stored insect pest products (Nawrot and Harmatha, 2012). Many re­
Saponin has shown harmful effects on Tribolium castaneum, Bufo viridis,
searchers have evaluated such compounds for insecticidal action against
and Lebistes reticulatus (Afolayan et al., 2014; Martin and Blossey,
major pests (Table 2) (Nawrot and Harmatha, 2012; Nozzolillo et al.,
2013; Sami et al., 2018). Several experiments also revealed that Quillaja
1997; Shany et al., 1970; Sylwia et al., 2006). Because of repellent or
Saponaria saponins cause fatality and intense cytotoxic activity in
deterrent activity, saponins directly affect insect pests’ reproduction and
certain insects such as Drosophila melanogaster (De Geyter et al., 2012;
growth (Singh and Kaur, 2018). These raise mortality rates by reducing
Smagghe et al., 2010). The Spodoptera littoralis Boisduval cotton leaf­
food consumption and disrupting the insect’s gut activity due to toxicity
worm has a broad host range and the key pest in many crops in different
and decreased digestibility (Adel et al., 2000). Saponins form complexes
parts of the world (De Geyter et al., 2007a). Saponin’s insecticidal action
with digestive enzymes such as proteases and thus appear to influence
against this species has been reported in many studies. The leek moth
the intestinal digestion process. Due to their permeability of the mem­
(Acrolepiopsis assectella) is an effective pest for cultivated plants such as
brane, saponins split the inner lining of the intestines mucosal cells.
leek, onion, and garlic. Bulb or leave tissues of host plants is the feed of
this pest (Mason et al., 2011). Poisonous compounds from A. porrum
Table 2 dried flowers and leaves were isolated, and toxic compounds’ adverse
Saponins Insecticidal activities in plants.
effects on leek moth growth and development were reported (Harmatha
Source Plant species Effect Reference et al., 1987; Mskhiladze et al., 2008; Sobolewska et al., 2016). The active
Leaf, shoot, Alfalfa cultivars Insecticidal to pea aphids Pedersen et al. compound was classified as a steroidal saponin, and its mortality levels
root (1976) (97%) in the diet recorded using flora. The protective function of soy­
Leaves, Trigonella Insecticidal to red flour Pemonge et al. beans (Glycine max L.) saponins against bruchid beetle (Callosobruchus
seeds foenumgraecum beetle and bean weevil (1997)
Chinensis L) stated by Applebaum et al. (1965). Larvae of beetles do not
Leaf Thevetia neriifolia Insecticidal to cotton Ray et al.
extracts leafworm (2013) hydrolyze saponins (Qasim et al., 2020). Szczepaniak et al. (2001) used
Bark Quillaja spp Insecticidal to cotton De Geyter et al. potato leaves that have been processed with various amounts of alfalfa
leafworm (2007b) saponins and used by larvae of Leptinotarsa decemlineata. L. decemlineata
Plant Quillaja saponaria Insecticidal to aphids De Geyter et al. has been the source for a substantial amount of potato losses and
(2012)
Flower, Pisum sativum Insecticidal to rice weevil Taylor et al.
decreased yield owing to its susceptibility to chemical pesticides. Larvae
seeds (2004) died after 4–6 days of leaves treated with alfalfa saponins (Szczepaniak
Leaves Pieris rapae Antifeedant Zhang et al. et al., 2001).
(2017)
Leaves Panax notoginseng Antifeedant, anti- Liu et al. (2018)
oviposition
4. Conclusion
Roots, C. graveolens Insecticidal to termite Rattan et al.
rhizomes (2015) Saponins have evident toxicity, as they work rapidly and robustly
Roots, Clematis graveolens Insecticidal to aphid, Rattan et al. against a wide range of pathogens. The most noticeable effects of sa­
rhizomes termites (2015)
ponins against pathogens were increased mortality and reduced food
Stem bark Catunaregam Antifeedant to DBM Gao et al.
spinosa (2011) intake, weight loss, reduced development, and reproduction activities.
Leaves B. vulgaris Antifeeding to DBM Shinoda et al. Several eco-friendly techniques have been used for stored grain products
(2002) and insect pests of field crops management. Besides, plant extracts from
Leaf Alfalfa plants Insecticidal to European Nozzolillo et al. different tissues, including leaf, stem, and flower, have been continu­
corn borer (1997)
ously used to manage pathogens. Consequently, the emphasis has been

24
M. Zaynab et al.
placed on characterizing and extracting plant metabolites from various
tissues. Such metabolites were eventually classified according to their
ability and behaviors in different groups. Many of the described com­
pounds had insecticidal, antifungal, and antibacterial effects at various
life stages of microorganisms, while other molecules had antimicrobial
effects. Such compounds were primarily insecticidal and antibacterial,
which were characterized as triterpenoid or steroidal saponins. Such
saponin compounds are significant in inhibiting the growth of micro­
organisms by disrupting specific enzymatic activity. The industrial
methods of separation and purification for saponins and the association
of saponin molecules must be strengthened. Knowledge of the molecular
mechanisms of a microbe immune system can also be useful. Such mo­
lecular pathways can be exploited by different molecular approaches (e.
g., CRISPR-Cas9 and RNAi) to modify microorganisms’ genetic makeup.
This effort will be of considerable assistance to the management of
pathogens.
Credit author statement
Madiha Zaynab, Yasir Sharif and Safdar Abbas write the manuscript.
Muhammad Zohaib Afzal, Muhammad Qasim7, Ahlam Khalofah, revised
the manuscript. Mohammad Javed Ansari and Li Tao, Khalid Ali Khan,
Shuangfei Li provide financial support and supervision.
Ethics
This manuscript work did not involves the use of human subjects.
Declaration of competing interest
The authors declare that there are no conflicts of interest.
Acknowledgment
Shenzhen science and technology application demonstration project
(KJYY20180201180253571) and special funds for science technology
innovation and industrial development of Shenzhen Dapeng New Dis­
trict(Grand No. KJYF202001-27). The authors extend their appreciation
to the Deanship of Scientific Research at King Khalid University for
funding this work through General Research Project under grant number
(R.G.P.1/192/41) and Research Center for Advanced Materials Science,
King Khalid University, Saudi Arabia for support.
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