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Defensive chemicals used by organisms for protection against potential consumers are generally products of secondary metabolism. Such chemicals are characteristic of free-living organisms with a limited range of movement or limited control over their movements. Despite the fact that chemical defense is widespread among animals as well as plants, the vast majority of theories advanced to account for patterns of allocation of energy and materials to defensive chemistry derive exclusively from studies of plant-herbivore interactions. Many such theories place an undue emphasis on primary physiological processes that are unique to plants (e.g., photosynthesis), rendering such theories limited in their utility or predictive power. The general failure of any single all-encompassing theory to gain acceptance to date may indicate that such a theory might not be a biologically realistic expectation. In lieu of refining theory, focusing attention on the genetic and biochemical mechanisms that underlie chemical defense allocation is likely to provide greater insights into understanding patterns across taxa. In particular, generalizations derived from understanding such mechanisms in natural systems have immediate applications in altering patterns of human use of natural and synthetic chemicals for pest control.

Transgenic tobacco (Nicotiana tabacum L.) plants overexpressing the enzyme L-phenylalanine ammonia-lyase (PAL; EC 4.3.1.5) were grown from seeds of a primary transformant containing the bean PAL2 gene, which had shown homology-dependent silencing of the endogenous tobacco PAL genes. Analysis of endogenous and transgene-encoded PAL transcripts and protein in the primary transformant (T0) and first-generation (T1) overexpressor plants indicated that the transgene-encoded PAL is the cause of the greater than wild-type levels of PAL activity (up to 5- and 2-fold greater in leaf and stem tissue, respectively) in the T1 plants. Leaves of PAL-overexpressing plants contained increased levels of the hydroxycinnamic acid ester chlorogenic acid but not of the flavonoid rutin, indicating that PAL is the key control point for flux into chlorogenic acid. In addition, levels of the glucoside of 4-coumaric acid increased in the overexpressing plants, suggesting that the 4-coumarate:coenzyme A ligase or coumarate hydroxylase reactions might have become limiting. These results help to define the regulatory architecture of the phenylpropanoid pathway and indicate the possibility of engineering-selective changes in this complex metabolic pathway by overexpression of a single early pathway gene

Phenylalanine ammonia-lyase (PAL) catalyzes the first step in phenylpropanoid synthesis. The role of PAL in pathway regulation was investigated by measurement of product accumulation as a function of enzyme activity in a collection of near-isogenic transgenic tobacco plants exhibiting a range of PAL levels from wild type to 0.2% of wild type. In leaf tissue, PAL level is the dominant factor regulating accumulation of the major product chlorogenic acid and overall flux into the pathway. In stems, PAL at wild-type levels contributes, together with downstream steps, in the regulation of lignin deposition and becomes the dominant, rate-determining step at levels 3- to 4-fold below wild type. The metabolic impact of elevated PAL levels was investigated in transgenic leaf callus that overexpressed PAL. Accumulation of the flavonoid rutin, the major product in wild-type callus, was not increased, but several other products accumulated to similarly high levels. These data indicate that PAL is a key step in the regulation of overall flux into the pathway and, hence, accumulation of major phenylpropanoid products, with the regulatory architecture of the pathway poised so that downstream steps control partitioning into different branch pathways.

Phenolics are one of the many secondary metabolites implicated in allelopathy. To establish that allelopathy functions in a natural ecosystem, the allelopathic bioassay must be ecologically realistic so that responses of appropriate bioassay species are determined at relevant concentrations. It is important to isolate, identify, and characterize phenolic compounds from the soil. However, since it is essentially impossible to simulate exact field conditions, experiments must be designed with conditions resembling those found in natural systems. It is argued that allelopathic potential of phenolics can be appreciated only when we have a good understanding of 1) species responses to phenolic allelochemicals, 2) methods for extraction and isolation of phenolic allelochemicals, and 3) how abiotic and biotic factors affect phenolic toxicity.

It has been proposed that natural products synthesized by plants contribute to their resistance to pests and pathogens. We show here that transgenic tobacco plants with suppressed levels of the phenylpropanoid biosynthetic enzyme phenylalanine ammonia-lyase (L-phenylalanine ammonia-lyase, EC 4.3.1.5) and correspondingly low levels of chlorogenic acid, the major soluble leaf phenylpropanoid product, exhibit more rapid and extensive lesion development than wild-type plants after infection by the virulent fungal pathogen Cercospora nicotianae. These observations provide direct evidence that phenylpropanoid products contribute to disease limitation. No induction of transcripts encoding phenylalanine ammonia-lyase or the lignin branch pathway enzyme caffeic acid O-methyltransferase was observed during the infection and there was no perturbation in the pattern of soluble phenylpropanoids. Hence, increased disease susceptibility does not involve inhibition of a pathogen-induced response but likely reflects inhibition of the developmental accumulation of chlorogenic acid. Demonstration of the contribution of such preformed protectants to plant health identifies attractive targets for manipulation by breeding or gene transfer to reduce the quantitative impact of disease

Allocation of allomones of transgenic Bacillus thuringiensis Gossypium hirsutum (Bt cotton) (cv. GK-12) and non-Bt-transgenic cotton (cv. Simian-3) grown in elevated CO2 in response to infestation by cotton aphid, Aphis gossypii Glover, was studied in a closed-dynamics CO2 chamber. Significant increases in foliar condensed tannin and carbon/nitrogen ratio for GK-12 and Simian-3 were observed in elevated CO2 relative to ambient CO2, as partially supported by the carbon nutrient balance hypothesis, owing to limiting nitrogen and excess carbon in cotton plants in response to elevated CO2. The CO2 level significantly influenced the foliar nutrients and allomones in the cotton plants. Aphid infestation significantly affected foliar nitrogen and allomone compounds in the cotton plants. Allomone allocation patterns in transgenic Bt cotton infested by A. gossypii may have broader implications across a range of plant and herbivorous insects as CO2 continues to rise.

The creation of artificial metabolic sinks in plants by genetic engineering of key branch points may have serious consequences for the metabolic pathways being modified. The introduction into potato of a gene encoding tryptophan decarboxylase (TDC) isolated from Catharanthus roseus drastically altered the balance of key substrate and product pools involved in the shikimate and phenylpropanoid pathways. Transgenic potato tubers expressing the TDC gene accumulated tryptamine, the immediate decarboxylation product of the TDC reaction. The redirection of tryptophan into tryptamine also resulted in a dramatic decrease in the levels of tryptophan, phenylalanine, and phenylalanine-derived phenolic compounds in transgenic tubers compared with nontransformed controls. In particular, wound-induced accumulation of chlorogenic acid, the major soluble phenolic ester in potato tubers, was found to be two- to threefold lower in transgenic tubers. Thus, the synthesis of polyphenolic compounds, such as lignin, was reduced due to the limited availability of phenolic monomers. Treatment of tuber discs with arachidonic acid, an elicitor of the defense response, led to a dramatic accumulation of soluble and cell wall-bound phenolics in tubers of untransformed potato plants but not in transgenic tubers. The transgenic tubers were also more susceptible to infection after inoculation with zoospores of Phytophthora infestans, which could be attributed to the modified cell wall of these plants. This study provides strong evidence that the synthesis and accumulation of phenolic compounds, including lignin, could be regulated by altering substrate availability through the introduction of a single gene outside the pathway involved in substrate supply. This study also indicates that phenolics, such as chlorogenic acid, play a critical role in defense responses of plants to fungal attack.

We conducted two experiments to determine how resources influenced the intraspecific and within-plant allocation by tomato plants (Lycopersicon esculentum) to the soluble phenolics rutin and chlorogenic acid. We also measured the effect of resource availability on growth by measuring mass and other physical and cellular attributes of the plant. In the first experiment, we subjected plants to four levels of potassium nitrate fertilizer. In the second experiment, we subjected plants to high and low levels of potassium nitrate fertilizer and light. Both experiments yielded results consistent with the growth-differentiation balance hypothesis. Plants grown with low resources showed low levels of soluble phenolics and low plant mass. Plants grown with intermediate levels of resources showed high phenolic concentrations but inhibited growth. Plants grown with high resources had high growth but no increases in phenolic concentrations. The results were also consistent with the general prediction of the optimal defense hypothesis that there should be a negative relationship between growth and defense. We discuss possible adaptive explanations for the specific patterns observed. There were also consistent within-plant differences in phenolic concentrations. These differences in phenolic concentrations were large enough to have potential consequences for insect herbivores feeding on tomato plants.

Allelopathy and resource competition have often been suggested to explain plant-plant interference. Many studies have attempted to separate these two mechanisms of interference to demonstrate either as a probable cause of an observed growth pattern. We, however, are of the opinion that separating allelopathy from resource competition is essentially impossible in natural systems. Furthermore, any experimental design to separate allelopathy and resource competition will create conditions that will never occur in nature. In this article, the ecological interaction between allelopathy and resource competition in natural systems is discussed.

Aqueous extracts of Ailanthus altissima bark and foliage were previously shown to be toxic to other plants. Using bioassay-directed fractionation, I isolated the phytotoxic compound from A. altissima root bark and identified it to be ailanthone, a quassinoid compound having molecular mass of 376. Ailanthone was highly phytotoxic, with concentrations of 0.7 ml/L causing 50% inhibition of radicle elongation in a standardized bioassay with garden cress (Lepidium sativum) seeds. Ailanthone exhibited potent pre- and postemergence herbicidal activity in greenhouse trials. Postemergence activity was especially striking; even the lowest application rate (0.5 kg/ha) caused complete mortality of five of the seven plant species tested within 5 d of treatment. In contrast, the highest application rate (8 kg/ha) did not cause any detectable injury to A. altissima seedlings, indicating the presence of a protective mechanism in the producer species to prevent autotoxicity. Ailanthone was rapidly detoxified in field soil as a result of microbial activity. Applications of ailanthone equivalent to 0.5 and 4.0 kg/ha completely lost their phytotoxicity within less than or equal to 5 d when incubated in the presence of nonsterile soil. When incubated with sterile soil under identical conditions, however, ailanthone remained highly phytotoxic throughout the 21-d duration of the investigation. The high level of postemergence herbicidal activity in conjunction with its rapid biodegradation in soil suggest ailanthone may have potential for development as a natural-product herbicide