Phenolic secondary metabolites play an important role in plant-derived food quality, as they affect quality characteristics such as appearance, flavour and health-promoting properties. Their content in foods is affected by many factors that influence phenolic stability, biosynthesis and degradation. In terms of their biosynthesis the key enzyme phenylalanine ammonia-lyase (PAL) is especially relevant, as it can be induced by different stress (environmental) conditions. In addition, polyphenol oxidases (PPO) and peroxidases (POD) are the main enzymes responsible for quality loss due to phenolic degradation. The different factors affecting phenolic-related food quality are reviewed. These include internal (genetic) and environmental (agronomic) factors, technological treatments applied during postharvest storage of fruits and vegetables, as well as processing and storage of the processed products. The different strategies that are required to either maintain or enhance the phenolic-related quality of foods are critically reviewed. Genetic modification designed to decrease polyphenol oxidases or peroxidases is not always a feasible method, owing to side problems related to the growth and defence of the plant. Agronomic treatments can be used to enhance the phenolic content and pigmentation of fruits and vegetables, although the information available on this topic is very scarce and even contradictory. Some postharvest treatments (cold storage, controlled or modified atmospheres, etc) can also improve phenolic-related quality, as well as new processing methods such as irradiation (gamma, UV), high-field electric pulses, high hydrostatic pressures and microwaves. Les composes phenoliques, metabolites secondaires des plantes, jouent un role dans la qualite des fruits, au niveau de l'apparence, de la flaveur et des qualites nutritives. L'enzyme cle de leur biosynthese est la phenylalanine ammonium-lyase, regulee par les facteurs environnementaux. La polyphenol oxydase et la peroxydase sont responsables des degradations donc des pertes de qualite. Les differents facteurs internes (genetiques) et externes (agronomiques) affectant la qualite des produits vegetaux sont etudies. Les differentes mesures envisageables (modification genetique, traitements chimiques ou physiques apres recolte) pour modifier le contenu en compose phenoliques et donc la qualite des produits vegetaux sont etudies.

Phenolic compounds and related enzymes as determinants of quality in fruits and vegetables

Lignin, one of the most abundant terrestrial biopolymers, is indispensable for plant structure and defense. With the availability of the full genome sequence, large collections of insertion mutants, and functional genomics tools, Arabidopsis constitutes an excellent model system to profoundly unravel the monolignol biosynthetic pathway. In a genome-wide bioinformatics survey of the Arabidopsis genome, 34 candidate genes were annotated that encode genes homologous to the 10 presently known enzymes of the monolignol biosynthesis pathway, nine of which have not been described before. By combining evolutionary analysis of these 10 gene families with in silico promoter analysis and expression data (from a reverse transcription-polymerase chain reaction analysis on an extensive tissue panel, mining of expressed sequence tags from publicly available resources, and assembling expression data from literature), 12 genes could be pinpointed as the most likely candidates for a role in vascular lignification. Furthermore, a possible novel link was detected between the presence of the AC regulatory promoter element and the biosynthesis of G lignin during vascular development. Together, these data describe the full complement of monolignol biosynthesis genes in Arabidopsis, provide a unified nomenclature, and serve as a basis for further functional studies.

http://www.plantphysiol.org/content/plantphysiol/133/3/1051.full.pdf

Genome-wide characterization of the lignification toolbox in Arabidopsis.

The flavonoid biosynthetic pathway in Arabidopsis: Structural and genetic diversity

Arabidopsis ecotype Columbia (Col-0) seedlings, transformed with a phenylalanine ammonia-lyase 1 promoter (PAL1)-[beta]-glucuronidase (GUS) reporter construct, were inoculated with virulent and avirulent isolates of Peronospora parasitica. The PAL1 promoter was constitutively active in the light in vascular tissue but was induced only in the vicinity of fungal structures in the incompatible interaction. A double-staining procedure was developed to distinguish between GUS activity and fungal structures. The PAL1 promoter was activated in cells undergoing lignification in the incompatible interaction in response to the pathogen. Pretreatment of the seedlings with 2-aminoindan-2-phosphonic acid (AIP), a highly specific PAL inhibitor, made the plants completely susceptible. Lignification was suppressed after AIP treatment, and surprisingly, pathogen-induced PAL1 promoter activity could not be detected. Treatment of the seedlings with 2-hydroxyphenylaminosulphinyl acetic acid (1,1-dimethyl ester) (OH-PAS), a cinnamyl alcohol dehydrogenase inhibitor specific for the lignification pathway, also caused a shift toward susceptibility, but the effect was not as pronounced as it was with AIP. Significantly, although OH-PAS suppressed pathogen-induced lignification, it did not suppress pathogen-induced PAL1 promoter activation. Salicylic acid (SA), supplied to AIP-treated plants, restored resistance and both pathogen-induced lignification and activation of the PAL1 promoter. Endogenous SA levels increased significantly in the incompatible but not in the compatible combination, and this increase was suppressed by AIP but not by OH-PAS. These results provide evidence of the central role of SA in genetically determined plant disease resistance and show that lignification per se, although providing a component of the resistance mechanism, is not the deciding factor between resistance and susceptibility.

/pdf/production-of-salicylic-acid-precursors-is-a-major-function-39qu0hl9as.pdf

Production of Salicylic Acid Precursors Is a Major Function of Phenylalanine Ammonia-Lyase in the Resistance of Arabidopsis to Peronospora parasitica

The phenylpropanoid pathway serves as a rich source of metabolites in plants, being required for the biosynthesis of lignin, and serving as a starting point for the production of many other important compounds, such as the flavonoids, coumarins, and lignans. In spite of the fact that the phenylpropanoids and their derivatives are sometimes classified as secondary metabolites, their relevance to plant survival has been made clear via the study of Arabidopsis and other plant species. As a model system, Arabidopsis has helped to elucidate many details of the phenylpropanoid pathway, its enzymes and intermediates, and the interconnectedness of the pathway with plant metabolism as a whole. These advances in our understanding have been made possible in large part by the relative ease with which mutations can be generated, identified, and studied in Arabidopsis. Herein, we provide an overview of the research progress that has been made in recent years, emphasizing both the genes (and gene families) associated with the phenylpropanoid pathway in Arabidopsis, and the end products that have contributed to the identification of many mutants deficient in the phenylpropanoid metabolism: the sinapate esters.

/pdf/the-phenylpropanoid-pathway-in-arabidopsis-l6bijxa925.pdf

The Phenylpropanoid Pathway in Arabidopsis

Phenylalanine ammonia-lyase (PAL) is encoded by a small family of genes in Arabidopsis. We cloned and partially characterized one of these genes, PAL1. The deduced amino acid sequence is highly similar to PAL from bean, parsley, and rice. The promoter contains sequence elements homologous to two putative regulatory elements conserved among several phenylpropanoid genes. The regulation of the PAL1 gene was examined by analysis of beta-glucuronidase (GUS) activity in transgenic Arabidopsis containing PAL1-GUS gene fusions. The PAL1 promoter was activated early in seedling development and in adult plants was strongly expressed in the vascular tissues of roots and leaves, but was not active in the root tip or the shoot apical meristem. In flowers, expression was observed in sepals, anthers, and carpels, but not in petals. Transcripts encoded by the endogenous PAL genes and GUS transcripts from the PAL1-GUS gene fusion were induced by wounding, HgCl2-stress, and light. Analysis of the regulatory properties of 5' deleted promoters showed that the proximal region of the promoter to -290 was sufficient to establish the full tissue-specific pattern of expression and that the proximal region to -540 was responsive to environmental stimuli. Negative and positive elements were located between -1816 and -823 and between -823 and -290, respectively.

Stephan Ohl

Papers

Functional properties of a phenylalanine ammonia-lyase promoter from Arabidopsis.