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Role of secondary metabolites in plant defense against pathogens
Accepted Manuscript Role of secondary metabolites in plant defense against pathogens Madiha Zaynab, Mahpara Fatima, Safdar Abbas, Yasir Sharif, Muhammad Umair, Muhammad Hammad Zafar, Khalida Bahadar PII:
S0882-4010(18)31278-6
DOI:
10.1016/j.micpath.2018.08.034
Reference:
YMPAT 3119
To appear in:
Microbial Pathogenesis
Received Date: 13 July 2018 Revised Date:
16 August 2018
Accepted Date: 18 August 2018
Please cite this article as: Zaynab M, Fatima M, Abbas S, Sharif Y, Umair M, Zafar MH, Bahadar K, Role of secondary metabolites in plant defense against pathogens, Microbial Pathogenesis (2018), doi: 10.1016/j.micpath.2018.08.034. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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Role of Secondary Metabolites in Plant Defense against Pathogens.
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Madiha Zaynab1*, Mahpara Fatima2*, Safdar Abbas*3 Yasir Sharif4, Muhammad Umair3,
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Muhammad Hammad Zafar4, Khalida Bahadar
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1. College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou 350002, PR.
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China.
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2. College of Crop Science, Fujian Agriculture and Forestry University, Fuzhou 350002, PR
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3. Department of Biochemistry, Faculty of Biological Sciences, Quaid-i-Azam University
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Islamabad, Pakistan.
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4. College of Plant Protection, Fujian Agriculture and Forestry University, Fuzhou 350002, PR.
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China.
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5. Faculty of Life science University of Agriculture Faisalabad
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6. PARC Institutes of Advanced Studies in Agriculture, NARC, Islamabad, Pakistan
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*Corresponding Author: MADIHA ZAYNAB ,
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[email protected]
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MAHPARA FATIMA [email protected]
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SAFDAR ABBAS [email protected]
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Abstract
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Pathogens get entry into host cell, reproduce there and use biological machinery of host
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plants which is threat to global crop production. Integrated management strategies based
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upon minimizing population and use of resistant cultivars can address this potential problem. In
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developing world farmers are less likely to adopt these approaches instead they prefer the use
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of chemical pesticides. Reckless use of chemical pesticides is destroying our ecosystem. That’s
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why it is required to explore ecofriendly alternatives, like plant based metabolites to control
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pathogens. Studies conducted on different plant-metabolites reported that these metabolite
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can potentially combat plant pathogens. In this study we have also discussed some of plant
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secondary metabolites including alkaloids, flavonoids and phenolics. In this review we tried to
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highlight the new trends in utilizing secondary metabolites for controlling bacterial, viral and
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fungal pathogens with the hope that upcoming drugs will be human and ecosystem friendly.
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Keywords: Plant protection, Natural Product, Phytochemicals, Food security, Immunity.
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Introduction
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Plants have evolved complex defense system to overcome the biotic and abiotic stresses as
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natural systems poses plenty of opposing forces on plants [1]. Variety of stress forces together
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affects the plants so, any change in metabolic physiology of plant cannot be referred to be
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associated with a specific particular stress factor. In context of specific stress, several response
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pathways are invoked, and in signaling response pathways for pathogens and herbivorous
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insects, several inter-connections exists [2-10]. Some of these response pathways are induced by
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infection and some are performed regardless of antimicrobial nature. Formation of pathogen’s
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cell wall degrading enzymes and synthesis of polymeric barriers to hinder pathogen entrance are
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some of the other means of plant defense [11]. Additionally plants have specific recognition and
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signaling systems that enables plant to detect pathogen entrance rapidly and initiate an effective
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defense response [12]. Plants have also evolved to respond subsequent microbes attack in case if
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they get infected.
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Plants synthesize diversity of secondary metabolites which prominently function to protect plants
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against predators and microbes according to toxic nature of microbes and repel the microbes and
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herbivores. Some secondary metabolites help plant to communicate with other organisms and
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some protects plants from abiotic stress e.g. UV-B radiations [13], so these secondary
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metabolites are significantly important for growth and development [14]. Three major types of
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secondary metabolites viz. Phenolics, Terpenes and Nitrogen/Sulfur containing compounds are
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produced in plant’s body. Terpenes have 5-C isoterpenoid as their basic unit that are toxins and
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deters herbivores. Shikimic acid pathway gives the products that forms phenolics which
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imparts defensive ability to plants. Nitrogen and sulfur containing compounds are mainly
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synthesized from amino acids [15, 16]. Defensive role of plant’s secondary metabolites have
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been confirmed by in vitro examining of plants for which expression of secondary metabolites
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was modified by modern techniques [17, 18]. Formation of secondary metabolites is the result of
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millions of years of plant’s interaction with pathogens and it is considered that more than
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100,000 metabolites are known to be involve in plant defense system, so the situation is still not
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clear [19]. Although it is considered that plant with high concentration of secondary metabolites
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is more resistant to biotic and abiotic stresses but their production is thought to be expensive for
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plant growth and reproduction [20, 21]. Function and structure of plant secondary
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metabolites explain that why plants have evolved induced defense, which is characterized
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with increased concentration in stress situations [22]. Several studies have uncovered that
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hundreds of plant compounds possess ecological and chemical defensive role, which have
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opened a new area of research known as ecological biochemistry [23, 24].
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Secondary metabolites in plant interactions with pathogens
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From long time, secondary metabolites have been suggested to interact with pathogenic
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organisms [25, 26] and among longest plant immune response studies one is involvement of
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secondary metabolites in plant interactions with pathogens (Fig.1) [27, 28]. Decades of research
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have demonstrated that a large number of secondary metabolites have proven their role in plant
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defense response to pathogens. Functions of secondary compounds are heavily pooled in
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conserved framework aside from their high structural diversity and several biosynthetic
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pathways are not conserved in plant kingdom [29]. Production and activation of these
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compounds like other conserved mechanisms of plant defense is facilitated by microbial
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detection via defense proteins or MAMPs recognition by pattern recognition patterns [30]. For
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the classification of secondary compounds several criteria has been introduced because of their
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diverse assortment in plant immunity. These criteria include common precursors, core structure
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and mechanism of action. Commonly used classification based on way of synthesis and
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accumulation of defense related phytochemicals. De novo production of metabolites due to an
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infection named as phytoalexins [28, 31] while, Phytoanticipins is a term used for production and
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storage of defense related metabolites in plant tissues [16].
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Secondary metabolites mode of action
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In plants defense system, widely distributed compounds are phenyl propanoids and flavonoids
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which possess different mode of action. Hundreds of antifungal drugs target only 6 processes,
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most of them act parallel to cell signaling compounds and effects physiological activities or act
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on parts of pathogen like; enzyme inhibition, DNA alkylation and reproductive system etc.[32].
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Mostly these compounds have hydroxyl group containing phenolics, which are likely to
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dissociate in phenolate ions. As phenolic hydroxyl groups form ionic bonds and hydrogen bonds
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with peptides and protons so, their higher number results in high astringency and denaturation
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[19]. Without confirmation i.e. proper three-dimensional structure, proteins cannot work
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properly. Protein properties are changed with any change in protein confirmation which can
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prevent crosstalk with other proteins and DNA/RNA. Secondary metabolites interact and
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changes three-dimensional structure of proteins by forming covalent bond with free SH., OH- or
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amino groups, resulting in loss of function or change in protein turnover. Polyphenols form
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hydrogen bonds and stronger ionic bonds. When these weak non-covalent bonds are formed
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concomitantly and act with a protein changes its flexibility resulting in inactivation of protein.
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Polar nature of phenols makes them less toxic compounds because their absorption is less after
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oral intake [19]. An experiment carried out to check the effect of resveratrol and pinosylvin
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(stilbene compounds) on fungi growth using wood decay test with stilbene impregnated aspen
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and birch samples. Plate experiment showed that at pinosylvin concentration which is enough to
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prevent fungal growth, resveratrol enhances the growth [33].
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Secondary metabolites against plant insect
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Secondary metabolites do not reduce the growth and development of plant instead they affect
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the fodder value of plant tissues where they are produce. These are either induced against the
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attack of microbes and insects (phytoalexins) or stored in inactive forms (phytoanticipins) (Fig.2)
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(Table 1). During herbivory, β-glucosidase activated phytoanticipins results in release of biocidal
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aglycones [34]. Hydrolyzation of glucosinolates by myrosinases during tissue disruption is
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classic example of phytoanticipins. Benzoxazinoids (BXs) are another example of
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phytoanticipins that are present among Poaceae. During tissue damage, their hydrolysis by
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plastid targeted β-glucosidase produces biocidal aglycone BXs that acts as insect repellent [35].
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On the other hand phytoalexins includes alkaloids, terpenoids, isoflavonoids, etc., that affects the
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performance of herbivores [36]. Aside from providing defense to plant, secondary metabolites
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improves the fitness of plant. Nuessly 2007 reported that maize HPR to Helecoverpa zea (corn
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eatworm) is due to C-glycosyl flavone maysine and chlorogenic acid. 4,4-dimethyl cyclooctene
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provides defense to sorghum against shoot fly [37].
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Secondary metabolites against plant viruses
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Several secondary compounds of plant metabolism like alkaloids, phenolics, and flavonoids play
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antiviral functions. Due to diversified structures alkaloids have many biologically active
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compounds that effects living organisms [38, 39]. Studies reported some 18000 alkaloids in
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ancient Chinese herbs with antiviral properties. Alkaloid 7-deoxy-trans-dihydronarciclasine
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discovered from plantain lilies (Hosta plantaginea) is anti TMV with least IC50 value i.e. 1.80
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µM [40]. Similarly Bruceine-D present in extract of Brassica javanica possess inhibitory effect
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against PVY, CMV and TMV [41]. It has been studied that application of 100 µgm/L Bruceine-
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D embedded extract of white goosefoot (Chenopodium amaranticolor) inhibited more than 90%
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PVY and CMV infection after 15 minutes [41]. Seventeen quassinoids with IC50 value of 3.42-
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5.66 have been identified as anti TMV infection [42]. Chen et al. [43] evaluated the anti TMV
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properties of Picarma quassioides wood extract and identified a quassinoid with positive results
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and several other β-carboline alkaloids. His further experiments suggested that combined
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application of β-carboline and quassinoids provide infection inhibition of 36.4%-68.4% as
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compared to 25%-47.4% infection inhibition by lonely application of β-carboline @ 50 µgm/l
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[43]. Similarly, An et al. [44] reported the anti TMV activities of 60 and 65% by application of
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Cynanchum
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demethoxytylophorine N-oxide) two alkaloids at 500 µgm/L concentration.
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Plant secondary metabolites as antifungal compounds
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Studies suggest that majority of secondary metabolites possesses antifungal characteristics [45].
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Allied phenolics and flavonoids constitute a large group of phytochemicals [46]. These
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compounds are present in fruit skins and leaves in high concentrations and take part in plant
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defense against pigmentation, UV resistance and disease resistance as shown in (Table 2) [47].
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Phenolics are known to change the cell permeability of microbes and also cause structural and
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functional deformation of membrane proteins which result in distraction of pH gradient, ATP
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production and conservation system, membrane bonded enzymes, substrate utilization for ATP
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production [48, 49].
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Antimicrobial compounds hinders the pathogens growth in apoplast. For example saponins with
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strong antifungal properties α-tomatine (tomato saponin) activates monomeric G-protein
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pathways and phosphotyrosine kinase which binds to cell membrane followed by cell component
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leakage which leads to ROS burst and Ca+2 elevation in Fusarium oxysporum [50]. Different
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saponins resistance against several pathogenic fungi are produced in different plant species [51].
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Large number of linear or cyclic and unsaturated or unsaturated isoprene units constitute the
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terpenes. Some of the best known terpenes are turpentine and camphor (metabolites of
extracts
containing
(7-demethoxytylophorine
and
7-
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odoriferous plants). Some of plant terpenes are industrial and medicinal importance e.g., taxol
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group [52]. Discovery of sesquiterpenoid phytoalexins, zealexins in F. graminearum infected
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maize was result of characterization of physiological responses. Zealexins showed resistance
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against A. flavus, F. graminearum and R. microspores [53]. Antioxidant, antitumor and anti-
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inflammatory properties of flavonoids have reported. Antibacterial activities of secondary
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metabolites have been suggested in one study. Flavonoids have also been reported in liquid
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extracts of legume leaves e.g. Zapoteca portoricensis. These studies suggest anti-pseudomonas
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aeruginosa activity secondary metabolites [54].
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Plant secondary metabolites as antibacterial compounds
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Analysis of Pseudomonas syringae infected Arabidopsis root exudates revealed the defensive
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role of antimicrobial compounds present in root exudates against this bacterium. Out of eight
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strains, seven are unable to cause infection and plants challenged with non-pathogenic strains
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produced more secondary metabolites. Antibacterial activity gained by non-pathogenic bacteria
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was moderate against non-infecting strains. Antibacterial activity of root exudation due to
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infecting strains was nonsignificant [55]. Studies have been carried to clearly demonstrate the
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role of resistive role of phytoanticipins and phytoalexins against pathogens growth. Brassica
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rapa is important world vegetable crop and its quality is affected by the disease caused by
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Xanthomonas campestris pv. campestris (Xcc). Phenolic compounds and glucosinolates can
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confer resistance to Brassica, but little work have done in this context [56].
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Future Prospects
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Plants developed defence system against several biotic and abiotic stresses with the passage
191
of time. Aside from the secondary metabolites either induced by infection or demonstrated,
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modern tools are required for correctly assessing the correlation between N and S application and
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resistance management in crops. Number of previous studies have revealed that N and S
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containing secondary metabolites production is affected by amount of these compounds in
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growth medium so, an optimum amount of these nutrients is required for the proper growth of
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plant and resistance against microbes and environmental stresses. Identification of SIR proteins
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induction mechanism might be a great achievement with minimized use of fungicides. For
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efficient organic farming SIR may become a vital tool. That’s why research for development of
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natural pesticides is needed now a days. Gene cassettes for complete metabolic pathways may be
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generated in long term and defense related secondary metabolites may be generated by metabolic
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engineering of plants or in bioreactors. This will help to quickly overcome the plant microbes or
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environmental stresses.
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Tabel 1. List of plant secondary metabolites against Insects. Listed secondary metabolites
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are shown their linked to a specific category and their target insect in specific plan Plants
Categories
Metabolites Terpenoids
Resistance against
Citrus
Terpenoid
Atta cephalotes
Pine
and Monoterpenes
fir Steroids
Common
Tobbaco
bark beetle
[57]
[58]
Phytoecdysones Insect
[59]
Trans-anethole
[60]
fern Terpenoids
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Limonene
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Secondary
thymol, litura
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and
Spodoptera
citronellal,
Phenolics
Wheat
Willow
Phenolics
Phenolics
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Phenolics
plant
Salix
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Benzoic acid
Strawberry
Benzoic acid
Cotton
padi Galerucella
[62]
lineola Operophtera
[63]
brumata Phenolics
Phenolics Phenolics
Rhopalosiphum [61]
Tetranychus
[64]
urticae Gossypol
Heliothis
[65]
virescens, Heliothis zea Alkaloids
Nightshade Alkaloid
Leptinotarsa
[66]
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potato Benzoxazinoides Gramineae
demissine
decemlineata
DIMBOA
Ostrinia
[67]
nubilalis Cassava
Cyrtomenus
CNglcs
bergi
Glucosides
Cyanogenic
Bitter
Amygdalin and Capnodis
Glucosides
almond
prunasin
tenebronis
Trifolium
Amygdalin and Hypera postica
Glucosides
repens
prunasin
Cyanogenic
Lotus
Cyanogenic
Zygaena
glucosides
filipendulae
CNglcs
Spodoptera
Cyanogenic
P.lunatus
Glucosides
eridania
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Table 2.List Phenolic compound role against Fungus Chemical
Fungus
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Cyanogenic
Glucosides
benzaldehyde
Botrytis cinerea
protocatechuic Colletotrichum acid
Reference [73] [74]
circinans
Salicylic acid
eutypa lata
[75]
Vanillic acid
Phytophthora
[76]
infestans Chlorogenic
Fusarium
acid
oxysporum
[50]
[69]
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[68]
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Cyanogenic
[70]
[71]
[73]
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Naringin
Penicillium
[77]
Aspergillus
[78]
Oleuropein
Phytophthora
[79]
Nobiletin
Phoma
[80]
tracheiphila Genistein
Monilinia
[81]
fructicola Hordatin A
Helminthosporium [82] sativum
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mechanism of species against microbial fungi and virus.
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Fig1.Schematic classification of Secondary metabolites that may be involved in defence
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Fig 2: Profiling Plant secondary metabolites and their role in plant defense system
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1.Plants synthesize diversity of secondary metabolites which prominently functions to protect plant.
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2.Secondary metabolites induced against the attack of microbes and insects 3.We have highlight the new trends in utilizing secondary metabolites for controlling bacterial,
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viral and fungal pathogens and insects.
S0882-4010(18)31278-6
DOI:
10.1016/j.micpath.2018.08.034
Reference:
YMPAT 3119
To appear in:
Microbial Pathogenesis
Received Date: 13 July 2018 Revised Date:
16 August 2018
Accepted Date: 18 August 2018
Please cite this article as: Zaynab M, Fatima M, Abbas S, Sharif Y, Umair M, Zafar MH, Bahadar K, Role of secondary metabolites in plant defense against pathogens, Microbial Pathogenesis (2018), doi: 10.1016/j.micpath.2018.08.034. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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Role of Secondary Metabolites in Plant Defense against Pathogens.
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Madiha Zaynab1*, Mahpara Fatima2*, Safdar Abbas*3 Yasir Sharif4, Muhammad Umair3,
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Muhammad Hammad Zafar4, Khalida Bahadar
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1. College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou 350002, PR.
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China.
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2. College of Crop Science, Fujian Agriculture and Forestry University, Fuzhou 350002, PR
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3. Department of Biochemistry, Faculty of Biological Sciences, Quaid-i-Azam University
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Islamabad, Pakistan.
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4. College of Plant Protection, Fujian Agriculture and Forestry University, Fuzhou 350002, PR.
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China.
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5. Faculty of Life science University of Agriculture Faisalabad
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6. PARC Institutes of Advanced Studies in Agriculture, NARC, Islamabad, Pakistan
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*Corresponding Author: MADIHA ZAYNAB ,
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[email protected]
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MAHPARA FATIMA [email protected]
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SAFDAR ABBAS [email protected]
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Abstract
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Pathogens get entry into host cell, reproduce there and use biological machinery of host
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plants which is threat to global crop production. Integrated management strategies based
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upon minimizing population and use of resistant cultivars can address this potential problem. In
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developing world farmers are less likely to adopt these approaches instead they prefer the use
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of chemical pesticides. Reckless use of chemical pesticides is destroying our ecosystem. That’s
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why it is required to explore ecofriendly alternatives, like plant based metabolites to control
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pathogens. Studies conducted on different plant-metabolites reported that these metabolite
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can potentially combat plant pathogens. In this study we have also discussed some of plant
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secondary metabolites including alkaloids, flavonoids and phenolics. In this review we tried to
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highlight the new trends in utilizing secondary metabolites for controlling bacterial, viral and
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fungal pathogens with the hope that upcoming drugs will be human and ecosystem friendly.
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Keywords: Plant protection, Natural Product, Phytochemicals, Food security, Immunity.
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Introduction
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Plants have evolved complex defense system to overcome the biotic and abiotic stresses as
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natural systems poses plenty of opposing forces on plants [1]. Variety of stress forces together
55
affects the plants so, any change in metabolic physiology of plant cannot be referred to be
56
associated with a specific particular stress factor. In context of specific stress, several response
57
pathways are invoked, and in signaling response pathways for pathogens and herbivorous
58
insects, several inter-connections exists [2-10]. Some of these response pathways are induced by
59
infection and some are performed regardless of antimicrobial nature. Formation of pathogen’s
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cell wall degrading enzymes and synthesis of polymeric barriers to hinder pathogen entrance are
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some of the other means of plant defense [11]. Additionally plants have specific recognition and
62
signaling systems that enables plant to detect pathogen entrance rapidly and initiate an effective
63
defense response [12]. Plants have also evolved to respond subsequent microbes attack in case if
64
they get infected.
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Plants synthesize diversity of secondary metabolites which prominently function to protect plants
66
against predators and microbes according to toxic nature of microbes and repel the microbes and
67
herbivores. Some secondary metabolites help plant to communicate with other organisms and
68
some protects plants from abiotic stress e.g. UV-B radiations [13], so these secondary
69
metabolites are significantly important for growth and development [14]. Three major types of
70
secondary metabolites viz. Phenolics, Terpenes and Nitrogen/Sulfur containing compounds are
71
produced in plant’s body. Terpenes have 5-C isoterpenoid as their basic unit that are toxins and
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deters herbivores. Shikimic acid pathway gives the products that forms phenolics which
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imparts defensive ability to plants. Nitrogen and sulfur containing compounds are mainly
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synthesized from amino acids [15, 16]. Defensive role of plant’s secondary metabolites have
75
been confirmed by in vitro examining of plants for which expression of secondary metabolites
76
was modified by modern techniques [17, 18]. Formation of secondary metabolites is the result of
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millions of years of plant’s interaction with pathogens and it is considered that more than
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100,000 metabolites are known to be involve in plant defense system, so the situation is still not
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clear [19]. Although it is considered that plant with high concentration of secondary metabolites
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is more resistant to biotic and abiotic stresses but their production is thought to be expensive for
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plant growth and reproduction [20, 21]. Function and structure of plant secondary
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metabolites explain that why plants have evolved induced defense, which is characterized
83
with increased concentration in stress situations [22]. Several studies have uncovered that
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hundreds of plant compounds possess ecological and chemical defensive role, which have
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opened a new area of research known as ecological biochemistry [23, 24].
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Secondary metabolites in plant interactions with pathogens
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From long time, secondary metabolites have been suggested to interact with pathogenic
88
organisms [25, 26] and among longest plant immune response studies one is involvement of
89
secondary metabolites in plant interactions with pathogens (Fig.1) [27, 28]. Decades of research
90
have demonstrated that a large number of secondary metabolites have proven their role in plant
91
defense response to pathogens. Functions of secondary compounds are heavily pooled in
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conserved framework aside from their high structural diversity and several biosynthetic
93
pathways are not conserved in plant kingdom [29]. Production and activation of these
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compounds like other conserved mechanisms of plant defense is facilitated by microbial
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detection via defense proteins or MAMPs recognition by pattern recognition patterns [30]. For
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the classification of secondary compounds several criteria has been introduced because of their
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diverse assortment in plant immunity. These criteria include common precursors, core structure
98
and mechanism of action. Commonly used classification based on way of synthesis and
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accumulation of defense related phytochemicals. De novo production of metabolites due to an
100
infection named as phytoalexins [28, 31] while, Phytoanticipins is a term used for production and
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storage of defense related metabolites in plant tissues [16].
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Secondary metabolites mode of action
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In plants defense system, widely distributed compounds are phenyl propanoids and flavonoids
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which possess different mode of action. Hundreds of antifungal drugs target only 6 processes,
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most of them act parallel to cell signaling compounds and effects physiological activities or act
106
on parts of pathogen like; enzyme inhibition, DNA alkylation and reproductive system etc.[32].
107
Mostly these compounds have hydroxyl group containing phenolics, which are likely to
108
dissociate in phenolate ions. As phenolic hydroxyl groups form ionic bonds and hydrogen bonds
109
with peptides and protons so, their higher number results in high astringency and denaturation
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[19]. Without confirmation i.e. proper three-dimensional structure, proteins cannot work
111
properly. Protein properties are changed with any change in protein confirmation which can
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prevent crosstalk with other proteins and DNA/RNA. Secondary metabolites interact and
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changes three-dimensional structure of proteins by forming covalent bond with free SH., OH- or
114
amino groups, resulting in loss of function or change in protein turnover. Polyphenols form
115
hydrogen bonds and stronger ionic bonds. When these weak non-covalent bonds are formed
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concomitantly and act with a protein changes its flexibility resulting in inactivation of protein.
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Polar nature of phenols makes them less toxic compounds because their absorption is less after
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oral intake [19]. An experiment carried out to check the effect of resveratrol and pinosylvin
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(stilbene compounds) on fungi growth using wood decay test with stilbene impregnated aspen
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and birch samples. Plate experiment showed that at pinosylvin concentration which is enough to
121
prevent fungal growth, resveratrol enhances the growth [33].
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Secondary metabolites against plant insect
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Secondary metabolites do not reduce the growth and development of plant instead they affect
124
the fodder value of plant tissues where they are produce. These are either induced against the
125
attack of microbes and insects (phytoalexins) or stored in inactive forms (phytoanticipins) (Fig.2)
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(Table 1). During herbivory, β-glucosidase activated phytoanticipins results in release of biocidal
127
aglycones [34]. Hydrolyzation of glucosinolates by myrosinases during tissue disruption is
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classic example of phytoanticipins. Benzoxazinoids (BXs) are another example of
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phytoanticipins that are present among Poaceae. During tissue damage, their hydrolysis by
130
plastid targeted β-glucosidase produces biocidal aglycone BXs that acts as insect repellent [35].
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On the other hand phytoalexins includes alkaloids, terpenoids, isoflavonoids, etc., that affects the
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performance of herbivores [36]. Aside from providing defense to plant, secondary metabolites
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improves the fitness of plant. Nuessly 2007 reported that maize HPR to Helecoverpa zea (corn
134
eatworm) is due to C-glycosyl flavone maysine and chlorogenic acid. 4,4-dimethyl cyclooctene
135
provides defense to sorghum against shoot fly [37].
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Secondary metabolites against plant viruses
137
Several secondary compounds of plant metabolism like alkaloids, phenolics, and flavonoids play
138
antiviral functions. Due to diversified structures alkaloids have many biologically active
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compounds that effects living organisms [38, 39]. Studies reported some 18000 alkaloids in
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ancient Chinese herbs with antiviral properties. Alkaloid 7-deoxy-trans-dihydronarciclasine
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discovered from plantain lilies (Hosta plantaginea) is anti TMV with least IC50 value i.e. 1.80
142
µM [40]. Similarly Bruceine-D present in extract of Brassica javanica possess inhibitory effect
143
against PVY, CMV and TMV [41]. It has been studied that application of 100 µgm/L Bruceine-
144
D embedded extract of white goosefoot (Chenopodium amaranticolor) inhibited more than 90%
145
PVY and CMV infection after 15 minutes [41]. Seventeen quassinoids with IC50 value of 3.42-
146
5.66 have been identified as anti TMV infection [42]. Chen et al. [43] evaluated the anti TMV
147
properties of Picarma quassioides wood extract and identified a quassinoid with positive results
148
and several other β-carboline alkaloids. His further experiments suggested that combined
149
application of β-carboline and quassinoids provide infection inhibition of 36.4%-68.4% as
150
compared to 25%-47.4% infection inhibition by lonely application of β-carboline @ 50 µgm/l
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[43]. Similarly, An et al. [44] reported the anti TMV activities of 60 and 65% by application of
152
Cynanchum
153
demethoxytylophorine N-oxide) two alkaloids at 500 µgm/L concentration.
154
Plant secondary metabolites as antifungal compounds
155
Studies suggest that majority of secondary metabolites possesses antifungal characteristics [45].
156
Allied phenolics and flavonoids constitute a large group of phytochemicals [46]. These
157
compounds are present in fruit skins and leaves in high concentrations and take part in plant
158
defense against pigmentation, UV resistance and disease resistance as shown in (Table 2) [47].
159
Phenolics are known to change the cell permeability of microbes and also cause structural and
160
functional deformation of membrane proteins which result in distraction of pH gradient, ATP
161
production and conservation system, membrane bonded enzymes, substrate utilization for ATP
162
production [48, 49].
163
Antimicrobial compounds hinders the pathogens growth in apoplast. For example saponins with
164
strong antifungal properties α-tomatine (tomato saponin) activates monomeric G-protein
165
pathways and phosphotyrosine kinase which binds to cell membrane followed by cell component
166
leakage which leads to ROS burst and Ca+2 elevation in Fusarium oxysporum [50]. Different
167
saponins resistance against several pathogenic fungi are produced in different plant species [51].
168
Large number of linear or cyclic and unsaturated or unsaturated isoprene units constitute the
169
terpenes. Some of the best known terpenes are turpentine and camphor (metabolites of
extracts
containing
(7-demethoxytylophorine
and
7-
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odoriferous plants). Some of plant terpenes are industrial and medicinal importance e.g., taxol
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group [52]. Discovery of sesquiterpenoid phytoalexins, zealexins in F. graminearum infected
172
maize was result of characterization of physiological responses. Zealexins showed resistance
173
against A. flavus, F. graminearum and R. microspores [53]. Antioxidant, antitumor and anti-
174
inflammatory properties of flavonoids have reported. Antibacterial activities of secondary
175
metabolites have been suggested in one study. Flavonoids have also been reported in liquid
176
extracts of legume leaves e.g. Zapoteca portoricensis. These studies suggest anti-pseudomonas
177
aeruginosa activity secondary metabolites [54].
178
Plant secondary metabolites as antibacterial compounds
179
Analysis of Pseudomonas syringae infected Arabidopsis root exudates revealed the defensive
180
role of antimicrobial compounds present in root exudates against this bacterium. Out of eight
181
strains, seven are unable to cause infection and plants challenged with non-pathogenic strains
182
produced more secondary metabolites. Antibacterial activity gained by non-pathogenic bacteria
183
was moderate against non-infecting strains. Antibacterial activity of root exudation due to
184
infecting strains was nonsignificant [55]. Studies have been carried to clearly demonstrate the
185
role of resistive role of phytoanticipins and phytoalexins against pathogens growth. Brassica
186
rapa is important world vegetable crop and its quality is affected by the disease caused by
187
Xanthomonas campestris pv. campestris (Xcc). Phenolic compounds and glucosinolates can
188
confer resistance to Brassica, but little work have done in this context [56].
189
Future Prospects
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Plants developed defence system against several biotic and abiotic stresses with the passage
191
of time. Aside from the secondary metabolites either induced by infection or demonstrated,
192
modern tools are required for correctly assessing the correlation between N and S application and
193
resistance management in crops. Number of previous studies have revealed that N and S
194
containing secondary metabolites production is affected by amount of these compounds in
195
growth medium so, an optimum amount of these nutrients is required for the proper growth of
196
plant and resistance against microbes and environmental stresses. Identification of SIR proteins
197
induction mechanism might be a great achievement with minimized use of fungicides. For
198
efficient organic farming SIR may become a vital tool. That’s why research for development of
199
natural pesticides is needed now a days. Gene cassettes for complete metabolic pathways may be
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generated in long term and defense related secondary metabolites may be generated by metabolic
201
engineering of plants or in bioreactors. This will help to quickly overcome the plant microbes or
202
environmental stresses.
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Tabel 1. List of plant secondary metabolites against Insects. Listed secondary metabolites
205
are shown their linked to a specific category and their target insect in specific plan Plants
Categories
Metabolites Terpenoids
Resistance against
Citrus
Terpenoid
Atta cephalotes
Pine
and Monoterpenes
fir Steroids
Common
Tobbaco
bark beetle
[57]
[58]
Phytoecdysones Insect
[59]
Trans-anethole
[60]
fern Terpenoids
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Limonene
Reference
SC
Secondary
thymol, litura
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and
Spodoptera
citronellal,
Phenolics
Wheat
Willow
Phenolics
Phenolics
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Phenolics
plant
Salix
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Benzoic acid
Strawberry
Benzoic acid
Cotton
padi Galerucella
[62]
lineola Operophtera
[63]
brumata Phenolics
Phenolics Phenolics
Rhopalosiphum [61]
Tetranychus
[64]
urticae Gossypol
Heliothis
[65]
virescens, Heliothis zea Alkaloids
Nightshade Alkaloid
Leptinotarsa
[66]
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potato Benzoxazinoides Gramineae
demissine
decemlineata
DIMBOA
Ostrinia
[67]
nubilalis Cassava
Cyrtomenus
CNglcs
bergi
Glucosides
Cyanogenic
Bitter
Amygdalin and Capnodis
Glucosides
almond
prunasin
tenebronis
Trifolium
Amygdalin and Hypera postica
Glucosides
repens
prunasin
Cyanogenic
Lotus
Cyanogenic
Zygaena
glucosides
filipendulae
CNglcs
Spodoptera
Cyanogenic
P.lunatus
Glucosides
eridania
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Table 2.List Phenolic compound role against Fungus Chemical
Fungus
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Cyanogenic
Glucosides
benzaldehyde
Botrytis cinerea
protocatechuic Colletotrichum acid
Reference [73] [74]
circinans
Salicylic acid
eutypa lata
[75]
Vanillic acid
Phytophthora
[76]
infestans Chlorogenic
Fusarium
acid
oxysporum
[50]
[69]
SC
plants
[68]
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Cyanogenic
[70]
[71]
[73]
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Naringin
Penicillium
[77]
Aspergillus
[78]
Oleuropein
Phytophthora
[79]
Nobiletin
Phoma
[80]
tracheiphila Genistein
Monilinia
[81]
fructicola Hordatin A
Helminthosporium [82] sativum
212
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Fig1.Schematic classification of Secondary metabolites that may be involved in defence
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Fig 2: Profiling Plant secondary metabolites and their role in plant defense system
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1.Plants synthesize diversity of secondary metabolites which prominently functions to protect plant.
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2.Secondary metabolites induced against the attack of microbes and insects 3.We have highlight the new trends in utilizing secondary metabolites for controlling bacterial,
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viral and fungal pathogens and insects.