Plants resistance mechanisms

Extracts of plants have been used as insecticides by humans since before the time of the Romans. Some of these extracts have yielded compounds useful as sources (e.g., pyrethrins, rotenoids, alkaloids), others as models (e.g., pyrethrins, physostigmine) of commercial insecticides. Recent technological advances which facilitate the isolation and identification of the bioactive constituents of plants should ensure the continued usefulness of plant compounds in commercial insect control, both as sources and models of new insect control agents and also as components in host plant resistance mechanisms. The focus in this paper will be on several classes of compounds, including limonoids, chromenes, ellagitannins, and methyl ketones, which were found to be components of the natural defenses of both wild and cultivated plants and which may be useful in commercial insect control. 

Plant furanocoumarins occur widely in nature and provide formidable obstacles to grazing by herbivorous animals. Some insect species have nevertheless adapted to circumvent this powerful host-plant-resistance mechanism. It has been proposed that the leaf-rolling habit of some insect species may be an evolutionary adaptation to avoid light and thus avoid the toxic effects of furanocoumarins (21). Also, evidence has recently been obtained that the capacity of at least one leaf-mining Insect species to detoxify furanocoumarins allows the utilization of furanocoumarln-contalnlng plants as hosts (29). 

They act as antipathogenic agents and thus affect the process of pathogenesis. They may act on the host through the Induction of plant resistance mechanisms such as stimulation of lignification or enhancement of phytoalexin production. (Please refer to the chapter by Salt and Kuc in this volume for further discussion of this type of compound.) They may act on the pathogen to accentuate elicitor release or to prevent infection (host penetration), colonization (inhibition of phytotoxin synthesis, extracellular enzyme production and action, or phytoalexin degradation) or reproduction. 

The increases in activities of both 1,3-3-glucan hydrolases and chitinase in response to infection appears to be part of a general plant resistance mechanism. The nature of the inducers of the enzymes is not clear but 3,6-3-glucan components of the cell walls of the invading fungi may be involved. This is supported by the observation that when melon seedlings were placed in a solution (0.08%) of the 3,6-3-glucan laminarin for 18h at 15 C there was a subsequent increase in the level of... 

In addition, naturally growing plants resist plant pathogen and Insect attack because resistance develops over time via natural selection (35). Also, most natural and crop plants have, as a part of their basic physical and chemical makeup, a wide array of mechanisms that help them resist pest attack. These Include chemical toxicants, repellents, altered plant nutrients, hairiness, thorns, and diverse combinations of these (35). 

CYP6D1 of the housefly (Musca domestica) has been found to hydroxylate cyper-methrin and thereby provide a resistance mechanism to this compound and other pyrethroids in this species (Scott et al. 1998 see also Chapter 12). Also, this insect P450 can metabolize plant toxins such as the linear furanocoumarins xanthotoxin and bergapten (Ma et al. 1994). This metabolic capability has been found in the lepi-dopteran Papilio polyxenes (black swallowtail), a species that feeds almost exclusively on plants containing furanocoumarins. 

Selective toxicity is also important in relation to the development of resistance or tolerance to pollutants from two distinct points of view. On the one hand, there is interest among scientists concerned with crop protection and disease control in mechanisms by which crop pests, vectors of disease, plant pathogens, and weeds develop resistance to pesticides. Understanding the mechanism should point to ways of overcoming resistance, for example, other compounds not affected by resistance mechanisms or synergists to inhibit enzymes that provide a resistance mechanism. On the other hand, the development of resistance can be a useful indication of the environmental impact of pollutants. 

As explained in Chapter 1, the toxicity of natural xenobiotics has exerted a selection pressure upon living organisms since very early in evolutionary history. There is abundant evidence of compounds produced by plants and animals that are toxic to species other than their own and which are nsed as chemical warfare agents (Chapter 1). Also, as we have seen, wild animals can develop resistance mechanisms to the toxic componnds prodnced by plants. In Anstralia, for example, some marsupials have developed resistance to natnrally occnrring toxins produced by the plants upon which they feed (see Chapter 1, Section 1.2.2). 

Plants are continually exposed to a vast array of potential phytopathogenic fungi nevertheless, plants resist to most of them by blocking fungal development soon after penetration. Resistance against pathogens can be distinguished in resistance at the species level (non-host resistance) and resistance at the cultivar level (race-cultivar resistance). Plants lack a circulatory system and antibodies and have evolved a defense mechanism that is distinct from the vertebrate immune... 

Our studies indicate that rapid metabolic detoxification of linear furanocoumarins is an effective resistance mechanism for K polyxenes against the toxic effects of these compounds. It has been postulated that the adaptation of some plants to produce angular furanocoumarins was in response to the reduced effectiveness of the linear furanocoumarins as deterrents for herbivores such as polyxenes (22). Such may Indeed be true, but our studies on the comparative detoxification of linear and angular furanocoumarins suggest that, at best, the presence of angular furanocoumarins in plants confers only a tenuous margin of relative "safety" against polyxenes. 

The effect of plant root exudation and exudation patterns on root colonization and expression of toxin production must be considered. For example, it may be important to determine the effect of root exudates from cold-stressed plants on these organisms, since the exudates apparently first appear just after the plants break winter dormancy (34). These data should provide information on root colonization potential, possible stimulation or reduction of toxin production, and mechanisms of plant resistance to the organisms. 

Three classes of plant-derived drugs, the vinca alkaloids (vincristine, vinblastine, and vinorelbine), the epipodo-phyllotoxins (etoposide and teniposide and the tax-anes (paclitaxel and taxotere), are used in cancer chemotherapy. These classes differ in their structures and mechanisms of action but share the multidrug resistance mechanism, since they are all substrates for the multidrug transporter P-glycoprotein. 

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