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BIOSYNTHESIS OF CONIFEROPHAGOUS BARK BEETLE PHEROMONES AND CONIFER ISOPRENOIDS: EVOLUTIONARY PERSPECTIVE AND SYNTHESIS

  • Steven J. Seybold (a1), Jörg Bohlmann (a2) and Kenneth F. Raffa (a3)

Abstract

In this overview we compare the significance and evolutionary history of two interacting biological systems, the conifer-feeding bark beetles (Coleoptera: Scolytidae) and their host conifers (Gymnospermae: Coniferales and Taxales). Isoprenoid natural products play key roles in the aggregation of the bark beetles and in the defence of the conifers. Our approach is to couple the most recent advances in the biochemical and molecular literature on these systems with ecological and behavioral data to compare monoterpenoid pheromone biosynthesis in scolytids with monoterpene biosynthesis in conifers. This synthesis reveals and evaluates the evolutionary redundancy occurring in the biochemical systems of the insect and host. Although host monoterpenes may be utilized directly or as derivatives in aggregation by scolytids, oxygenated monoterpenes that are behaviorally active for scolytids have been rarely identified from their coniferous hosts. De novo monoterpenoid biosynthesis in the Scolytidae, a process that is likely to be rare among metazoans, is substantially different from monoterpene biosynthesis in the conifers. The pathways appear to be shared only at the late-stage reactions that follow the formation of isopentenyl diphosphate. Little is known of the regulation of monoterpene biosynthesis in conifers, but scolytids positively regulate monoterpenoid biosynthesis using a sesquiterpenoid hormone, juvenile hormone, which does not occur in conifers. Little is known of the subcellular site of synthesis of monoterpenoids in scolytids, but conifer monoterpene biosynthesis is compartmentalized in the plastids, which do not occur in scolytid cells. In addition to bark beetles and conifers, the vertebrate model presents one of the few systems in which isoprenoid synthesis has been studied enough to provide a meaningful comparison. Possible unique features of monoterpenoid pheromone biosynthesis in scolytids relative to isoprenoid biosynthesis in vertebrates include the following: (1) a monoterpenoid end product; (2) a hypothetically scolytid-specific prenyl transferase (= geranyl diphosphate synthase) that catalyzes the condensation of two five-carbon (C5) units, but does not catalyze additional condensation reactions with the C5 monomelic unit; (3) a scolytid-specific monoterpene (myrcene) synthase; and (4) a scolytid-specific, transcriptional-level sesquiterpenoid isoprenoid regulatory mechanism. Features 2 and 3 may be shared with conifers. This review also updates the 1985 landmark scientific paper by John Borden by listing the references and species of coniferophagous Scolytidae for which aggregation pheromones have been identified since 1985.

Dans cette révision, nous comparons l’importance et l’évolution de deux systèmes biologiques interactifs, les scolytes (Coleoptera : Scolytidae) qui se nourrissent à même les conifères et les conifères (Gymnospermae : Coniferales et Taxales) qui leur servent d’hôtes. Les produits naturels isoprénoïdes jouent un rôle important dans les agrégations de scolytes et la défense des conifères. Nous avons choisi comme approche de combiner les dernières découvertes en biochimie et en biologie moléculaire sur ces systèmes avec des données écologiques et comportementales, de façon à comparer la biosynthèse des phéromones monoterpénoïdes chez les scolytes et la biosynthèse des monoterpènes des conifères. Cette synthèse démontre et évalue la redondance évolutive qui prévaut dans les systèmes biochimiques des insectes et de leurs hôtes. Bien que les monoterpènes puissent être utilisés par les scolytes directement ou comme dérivés dans les agrégations, les monoterpènes oxygénés des conifères qui élicitent une réponse comportementale de la part des scolytes ont rarement été identifiés. La biosynthèse de novo des monoterpénoïdes chez les Scolytidae, un processus qui est probablement rare chez les métazoaires, diffère substantiellement de la biosynthèse des monoterpènes chez les conifères. Les voies empruntées semblent les mêmes seulement au cours des derniers stades qui suivent la formation des diphosphates d’isopentényle. On connaît peu de choses au sujet de la régulation de la biosynthèse des monoterpènes chez les conifères, mais les scolytes contrôlent certainement la biosynthèse des monoterpénoïdes par l’intermédiaire d’une hormone sesquiterpénoïde, l’hormone juvénile, qui n’existe pas chez les conifères. On ne connaît pas très bien non plus le site de synthèse des monoterpénoïdes chez les scolytes, mais la biosynthèse des monoterpènes des conifères est compartimentée dans les plastes, que l’on ne retrouve pas dans les cellules des scolytes. En plus des scolytes et des conifères, le modèle vertébré possède l’un des rares systèmes dans lequel la synthèse des isoprénoïdes a été suffisamment étudiée pour permettre une comparaison intéressante. Parmi les caractéristiques possiblement exclusives à la biosynthèse des phéromones monoterpénoïdes chez les scolytes par comparaison à la biosynthèse des isoprénoïdes chez les vertébrés, il faut mentionner (1) l’obtention d’un produit final monoterpénoïde, (2) une prényl-transférase (= gényl-diphosphate synthase), hypothétiquement spécifique aux Scolytidae, qui catalyse la condensation de deux unités 5-carbone (C5), mais ne catalyse pas les réactions additionnelles de condensation avec l’unité monomère C5, (3) une monoterpène (myrcène) synthase spécifique aux Scolytidae et (4) un mécanisme régulateur du niveau de transcription sesquiterpénoïde-isoprénoïde qui soit spécifique aux scolytes. Les caratéristiques 2 et 3 se rencontrent peut-être aussi chez les conifères. Cette révision met à jour l’ouvrage classique de John Borden (Borden 1985) en présentant la liste des références et en énumérant les espèces de Scolytidae coniférophages chez lesquelles les phéromones d’agrégation ont été identifiées depuis 1985.

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1 Author to whom all correspondence should be addressed (E-mail: sseybokd@tc.umn.edu).

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