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Role of Secondary Metabolites in Defense Mechanisms of Plants Review Article Biology and Medicine, 3 (2) Special Issue: 232-249, 2011 eISSN: 09748369, www.biolmedonline.com Role of secondary metabolites in defense mechanisms of plants *Mazid M1, Khan TA2, Mohammad F1 1 Plant Physiology Division, Department of Botany, Faculty of Life Sciences, AMU, Aligarh, India. 2 Department of Biochemistry, Faculty of Life Sciences, AMU, Aligarh, India. *Corresponding Author: [email protected] Abstract In all natural habitats, plants are surrounded by an enormous number of potential enemies (biotic) and various kinds of abiotic environmental stress. Nearly all ecosystems contain a wide variety of bacteria, viruses, fungi, nematodes, mites, insects, mammals and other herbivorous animals, greatly responsible for heavy reduction in crop productivity. By their nature, plants protect themselves by producing some compounds called as secondary metabolites. Secondary metabolites, including terpenes, phenolics and nitrogen (N) and sulphur (S) containing compounds, defend plants against a variety of herbivores and pathogenic microorganisms as well as various kinds of abiotic stresses. This review presents an overview about some of the mechanisms by which plants protect themselves against herbivory, pathogenic microbes and various abiotic stresses as well as specific plant responses to pathogen attack, the genetic control of host-pathogen interactions. Keywords: Secondary metabolites; defense mechanism; phytoalexins; terpenes; alkaloids; phenolics; cynogenic glucosides. Abbreviations: GST, glucosinolate synthase transferase; SIR, systematic induced resistance; GSL, glucosinolates; GSH, glutathione; HCN, hydrogen cyanide. Introduction (Schaller et al., 1996). Once infected, some In natural systems, plants face a plethora of plants also develop immunity to subsequent antagonists and thus posses a myriad of microbial attacks (Putnam and Heisey, 1983; defense and have evolved multiple defense Putnam and Tang, 1986; Bernays, 1989; mechanisms by which they are able to cope Elakovich, 1987). with various kinds of biotic and abiotic stress Plants produce a high diversity of (Ballhorn et al., 2009). The most significant natural products or secondary metabolites with biotic and abiotic and man-made stress factors a prominent function in the protection against are summarized in figure 1. Generally, it is predators and microbial pathogens on the difficult to assign a change in the physiology of basis of their toxic nature and repellence to metabolism of the crop to a specific stress herbivores and microbes and some of which factor as normally a complex variety of various also involved in defense against abiotic stress stress factors affects the plant simultaneously. (e.g. UV-B exposure) and also important for However, there are inter-connections that exist the communication of the plants with other between distinct and opposing signaling organisms (Schafer et al., 2009), and are response pathways for defense against insignificant for growth and developmental pathogens and insect herbivores and there processes (Rosenthal et al., 1991). There are also appear to be multiple response pathways three major groups of secondary metabolites invoked, depending on the specific stress viz terpenes, phenolics and N and S context (Agosta, 1996; Barbour et al., 1987; containing compounds. Terpenes composed of Bostock, 1999; Bostock et al., 2001; Hell, 5-C isopentanoid units, are toxins and feeding 1997; Thomma et al., 1998; Vijayan et al., deterrents to many herbivores. Phenolics 1998; Whittaker and Feeny, 1971; Kusnierczyk synthesized primarily from products of the et al., 2007). Besides antimicrobial nature, shikimic acid pathway, have several important some of which are performed and some of defensive role in the plants. Members of the which induced by infection. There are various third major group i.e. N and S containing other modes of defense include the compounds are synthesized principally from construction of polymeric barriers to pathogen common amino acids (Rosenthal et al., 1992; penetration and the synthesis of enzymes that Van Etten et al., 2001). Recent in vitro degrade pathogen cell wall (Hammond et al., experiments using plants whose secondary 1996). In addition, plants employ specific metabolites expression has been altered by recognition and signaling systems enabling the modern molecular methods have to confirm rapid detection of pathogen invasion and their defensive roles (Mes et al., 2000; initiation of vigorous defensive responses Mansfield, 2000). Although the situation is still 232 MAASCON-1 (Oct 23-24, 2010): “Frontiers in Life Sciences: Basic and Applied” Review Article Biology and Medicine, 3 (2) Special Issue: 232-249, 2011 unclear, it is believed that most of the 100,000 Principal groups known secondary metabolites are to be Plant secondary metabolites can be divided involved in plant chemical defense systems, into three chemically distinct groups viz: which are formed throughout the millions of Terpenes, Phenolics, N and S containing years during which plants have co-existed with compounds. their attackers (Wink, 1999). Although higher concentrations of secondary metabolites might (i) Terpenes result in a more resistant plant, the production Terpenes constitute the largest class of of secondary metabolites is thought to be secondary metabolites and are united by their costly and reduces plant growth and common biosynthetic origin from acetyl-coA or reproduction (Simms, 1992; Karban and glycolytic intermediates (Gerhenzon et al., Baldwin, 1999; Simens et al., 2002). The cost 1991; Grayson, 1998; Fraga, 1988; Croteas, of defense has also been invoked to explain 1988; Loomis and Croteas, 1980; Robinson, why plants have evolved induced defense, 1980). A vast majority of the different terpenes where concentrations generally increase only structures produced by plants as secondary in stress situations (Harvell and Tollrian, metabolites that are presumed to be involved 1999). in defense as toxins and feeding deterrents to During the last several years, it has a large number of plant feeding insects and been discovered that hundreds of compounds mammals (Gershenzon and Croteau, 1991). that plants make have significant ecological Below, several examples will draw from the 5 and chemical defensive roles, opening a new major subclasses: area of scientific endeavour, often called ecological biochemistry (Harborne, 1988, (a) Monoterpenes (C10): Many derivatives are 1989). important agents of insect toxicity. For example, the pyrethroids (monoterpenes Secondary metabolites esters) occur in the leaves and flowers of Plants produce a large and diverse array of Chrysanthemum species show strong organic compounds that appear to have no insecticidal responses (neurotoxin) to insects direct functions in growth and development i.e. like beetles, wasps, moths, bees, etc. and a they have no generally recognised roles in the popular ingredient in commercial insecticides process of photosynthesis, respiration, solute because of low persistence in the environment transport, translocation, nutrient assimilation and low mammalian toxicity (Turlings et al., and differentiation (Hartmann, 1991). They 1995). have a very restricted distribution than primary In Gymnosperms (conifers) like Pine metabolites in the whole plant kingdom i.e. and Fir, monoterpenes accumulate in resin they are often found only in one plant species ducts found in the needles, twings and trunks or a taxonomically related group of species. mainly as α-pinene, β-pinene, limonene and High concentrations of secondary metabolites myrecene, all are toxic to numerous insects might result in a more resistant plant. Their including bark beetles, serious pest of conifer production is thought to be costly and reduces species throughout the world (Turlings et al., plant growth and reproduction (Simms, 1992; 1995). Karban and Baldwin, 1997; Harvell and Tollrian, 1999; Stotz et al., 1999; Siemens et (b) Sesquiterpenes (C15): A number of al., 2002). Therefore, defense metabolites can sesquiterpenes have been till now reported for be divided in to constitutive substances, also their role in plant defense such as costunolides called prohibitins or phytoanticipins and are antiherbivore agents of family composite induced metabolites formed in response to an characterized by a five membered lactone ring infection involving de novo enzyme synthesis, (a cyclic ester) and have strong feeding known as phytoalexins (Van Etten et al., 1994; repellence to many herbivorous insects and Grayer and Harborne, 1994). Phytoanticipins mammals (Picman, 1986). are high energy and carbon consuming and ABA is also a sesquiterpene plays exhibit fitness cost under natural conditions primarily regulatory roles in the initiation and (Mauricio, 1998), but recognized as the first maintenance of seed and bud dormancy and line of chemical defense that potential plants response to water stress by modifying pathogens have to overcome. In contrast, the membrane properties (Van-steveninck, phytoalexin production may take two or three 1983) and act as a transcriptional activator days, as by definition first the enzyme system (McCarty et al., 1991; Giraudat et al., 1992). In needs to be synthesized (Grayer and addition, it increases the cytosolic calcium Harborne, 1994). concentration and causes alkalinisation of the cytosol (Irving et al., 1992; Blatt and 233 MAASCON-1 (Oct 23-24, 2010): “Frontiers in Life Sciences: Basic and Applied” Review Article Biology and Medicine, 3 (2) Special Issue: 232-249, 2011
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