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      Plant Glandular Trichomes as Targets for Breeding or Engineering of Resistance to Herbivores

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          Abstract

          Glandular trichomes are specialized hairs found on the surface of about 30% of all vascular plants and are responsible for a significant portion of a plant’s secondary chemistry. Glandular trichomes are an important source of essential oils, i.e., natural fragrances or products that can be used by the pharmaceutical industry, although many of these substances have evolved to provide the plant with protection against herbivores and pathogens. The storage compartment of glandular trichomes usually is located on the tip of the hair and is part of the glandular cell, or cells, which are metabolically active. Trichomes and their exudates can be harvested relatively easily, and this has permitted a detailed study of their metabolites, as well as the genes and proteins responsible for them. This knowledge now assists classical breeding programs, as well as targeted genetic engineering, aimed to optimize trichome density and physiology to facilitate customization of essential oil production or to tune biocide activity to enhance crop protection. We will provide an overview of the metabolic diversity found within plant glandular trichomes, with the emphasis on those of the Solanaceae, and of the tools available to manipulate their activities for enhancing the plant’s resistance to pests.

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          Structure and function of enzymes involved in the biosynthesis of phenylpropanoids.

          As a major component of plant specialized metabolism, phenylpropanoid biosynthetic pathways provide anthocyanins for pigmentation, flavonoids such as flavones for protection against UV photodamage, various flavonoid and isoflavonoid inducers of Rhizobium nodulation genes, polymeric lignin for structural support and assorted antimicrobial phytoalexins. As constituents of plant-rich diets and an assortment of herbal medicinal agents, the phenylpropanoids exhibit measurable cancer chemopreventive, antimitotic, estrogenic, antimalarial, antioxidant and antiasthmatic activities. The health benefits of consuming red wine, which contains significant amounts of 3,4',5-trihydroxystilbene (resveratrol) and other phenylpropanoids, highlight the increasing awareness in the medical community and the public at large as to the potential dietary importance of these plant derived compounds. As recently as a decade ago, little was known about the three-dimensional structure of the enzymes involved in these highly branched biosynthetic pathways. Ten years ago, we initiated X-ray crystallographic analyses of key enzymes of this pathway, complemented by biochemical and enzyme engineering studies. We first investigated chalcone synthase (CHS), the entry point of the flavonoid pathway, and its close relative stilbene synthase (STS). Work soon followed on the O-methyl transferases (OMTs) involved in modifications of chalcone, isoflavonoids and metabolic precursors of lignin. More recently, our groups and others have extended the range of phenylpropanoid pathway structural investigations to include the upstream enzymes responsible for the initial recruitment of phenylalanine and tyrosine, as well as a number of reductases, acyltransferases and ancillary tailoring enzymes of phenylpropanoid-derived metabolites. These structure-function studies collectively provide a comprehensive view of an important aspect of phenylpropanoid metabolism. More specifically, these atomic resolution insights into the architecture and mechanistic underpinnings of phenylpropanoid metabolizing enzymes contribute to our understanding of the emergence and on-going evolution of specialized phenylpropanoid products, and underscore the molecular basis of metabolic biodiversity at the chemical level. Finally, the detailed knowledge of the structure, function and evolution of these enzymes of specialized metabolism provide a set of experimental templates for the enzyme and metabolic engineering of production platforms for diverse novel compounds with desirable dietary and medicinal properties.
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            Terpene synthases and the regulation, diversity and biological roles of terpene metabolism.

            Terpene synthases are the primary enzymes in the formation of low-molecular-weight terpene metabolites. Rapid progress in the biochemical and molecular analysis of terpene synthases has allowed significant investigations of their evolution, structural and mechanistic properties, and regulation. The organization of terpene synthases in large gene families, their characteristic ability to form multiple products, and their spatial and temporal regulation during development and in response to biotic and abiotic factors contribute to the time-variable formation of a diverse group of terpene metabolites. The structural diversity and complexity of terpenes generates an enormous potential for mediating plant-environment interactions. Engineering the activities of terpene synthases provides opportunities for detailed functional evaluations of terpene metabolites in planta.
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              Elucidation of the methylerythritol phosphate pathway for isoprenoid biosynthesis in bacteria and plastids. A metabolic milestone achieved through genomics.

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                Author and article information

                Journal
                Int J Mol Sci
                Int J Mol Sci
                ijms
                International Journal of Molecular Sciences
                Molecular Diversity Preservation International (MDPI)
                1422-0067
                2012
                12 December 2012
                : 13
                : 12
                : 17077-17103
                Affiliations
                [1 ]Department of Population Biology, Institute for Biodiversity and Ecosystem Dynamics, 1098 XH Science Park 904, Amsterdam, The Netherlands; E-Mails: j.j.glas@ 123456uva.nl (J.J.G.); b.c.j.schimmel@ 123456uva.nl (B.C.J.S.); j.m.albacano@ 123456uva.nl (J.M.A.)
                [2 ]Department of Plant Breeding, Subtropical and Mediterranean Horticulture Institute “La Mayora” (IHSM), Spanish Council for Scientific Research (CSIC), Experimental Station “La Mayora”, E-29750, Algarrobo-Costa, Málaga, Spain; E-Mail: rocio.escobar@ 123456eelm.csic.es
                [3 ]Department of Plant Physiology, Swammerdam Institute of Life Sciences, 1098 XH, Science Park 904, Amsterdam, The Netherlands; E-Mail: r.c.schuurink@ 123456uva.nl
                Author notes
                [* ]Author to whom correspondence should be addressed; E-Mail: m.kant@ 123456uva.nl ; Tel.: +31-20-5257-793; Fax: +31-20-5257-754.
                Article
                ijms-13-17077
                10.3390/ijms131217077
                3546740
                23235331
                98485068-1583-4762-9613-09f832a893de
                © 2012 by the authors; licensee Molecular Diversity Preservation International, Basel, Switzerland.

                This article is an open-access article distributed under the terms and conditions of the Creative Commons Attribution license ( http://creativecommons.org/licenses/by/3.0/).

                History
                : 06 November 2012
                : 28 November 2012
                : 05 December 2012
                Categories
                Review

                Molecular biology
                glandular trichome,plant-herbivore interactions,pathogen,solanaceae,pest resistance,plant defense,leaf hair,tomato,plant breeding,genetic engineering

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