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Variations in Allelochemical Composition of Leachates Of Variations in Allelochemical Composition of Leachates of Different Organs and Maturity Stages of Pinus halepensis Catherine Fernandez, Yogan Monnier, Elena Ormeño, Virginie Baldy, Stephane Greff, Vanina Pasqualini, Jean-Philippe Mévy, Anne Bousquet-Mélou To cite this version: Catherine Fernandez, Yogan Monnier, Elena Ormeño, Virginie Baldy, Stephane Greff, et al.. Varia- tions in Allelochemical Composition of Leachates of Different Organs and Maturity Stages of Pinus halepensis. Journal of Chemical Ecology, Springer Verlag, 2009, 35 (8), pp.970-979. 10.1007/s10886- 009-9667-8. hal-00603381v2 HAL Id: hal-00603381 https://hal-amu.archives-ouvertes.fr/hal-00603381v2 Submitted on 10 Apr 2018 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Variations in Allelochemical Composition of Leachates Pinus halepensis of Different Organs and Maturity Stages of Catherine Fernandez & Yogan Monnier & Elena Ormeño & Virginie Baldy & Stéphane Greff & Vanina Pasqualini & Jean-Philippe Mévy & Anne Bousquet-Mélou Abstract We investigated changes in the occurrence of showed that needles and, to a lesser extent, old roots, had allelochemicals from leachates of different Pinus halepensis higher chemical diversity than the roots of young and medium- organs taking into account the stages of pine stand age (i.e., aged pines. The highest diversity in phenolic constituents and young <15-years-old, middle age±30-years, and old > fatty acids was in young needles (Dchem=2.38). Finally, 60-years-old). GC-MS analysis of aqueous extracts caffeic acid, a compound that has allelopathic properties was revealed approx. 59 components from needles and roots. found in aqueous extracts at high concentrations in both The major constituents were divided into different phyto- young needles and old roots. The role of this compound in chemical groups—phenolics (50%), fatty acids (44%), and mediation of biological interactions in P. halepensis ecosystem terpenoids. Further analyses were carried out to characterize functioning is discussed. the distribution of allelochemicals in different organs and P. halepensis successional stages. Roots and needles had two Keywords Pinus halepensis Miller . Allepo pine . distinct chemical profiles, while needle leachates were Phenolic compounds . Terpenoids . Needles . Roots . Soil . composed mainly of oxygenated terpenoids (e.g., α- Allelopathy eudesmol, α-cadinol, and α-terpineol). Roots mainly contained fatty acids. Needles from young pine stands had the highest content of monoterpenes, suggesting their role as Introduction potential allelochemicals that could help young pine stands to establish. Pooling the different functional chemical groups Plant secondary metabolites affects ecosystem processes and biodiversity. Release of secondary metabolites into the environment, an important driver of biotic interactions, * : : : : C. Fernandez: ( ) Y. Monnier V. Baldy S. Greff occurs through litter decomposition, root exudates, vapor- J.-P. Mévy A. Bousquet-Mélou ization into the air, and leaching from plant parts to the soil Institut Méditerranéen d’Ecologie et Paléoécologie (IMEP)-UMR CNRS 6116, Equipe DFCV, Aix-Marseille Université, (Rice 1984). Production of secondary compounds is often Centre de St. Charles, Case 4, associated with plant protection. Compounds in above- 13331 Marseille Cedex 03, France ground parts help to protect plants against microbes, e-mail: [email protected] herbivores (Vernenghi et al. 1986), and/or UV irradiation V. Pasqualini (Delfine et al. 2003). Root compounds may be produced in Sciences Pour l’Environnement—UMR CNRS 6134, response to soil-borne pathogens (Valette et al. 1998). Equipe Feux, Université de Corse, Allelochemicals also may be involved with beneficial Faculté des Sciences et Techniques, interactions (Dicke et al. 2003), such as attracting polli- BP 52, 20250 Corte, France nators to flowers and leaves (Caissard et al. 2004)or E. Ormeño signaling events in plant-plant, plant-microbe, or plant- Division of Ecosystem Sciences, nematode interactions (Hiltpold and Turlings 2008). Department of Environmental Science, Policy, Among vascular plants, almost all allelochemicals are and Management (ESPM), Berkeley University of California, secondary metabolites and have the potential to impact 251A Mulford Hall Berkeley, CA 94720, USA ecosystem structure and function. Allelopathic components also play a role in regulating plant diversity (Chou 1999), (>60-years-old) and called successional stage “O”— establishment of invasive species (Bousquet-Mélou et al. mature forest with well-developed understory. 2005), and the dynamics in arid environments (Karageorgou Three replicates were collected from each successional et al. 2002). stage. Sites were selected along the Southern hillside of the In the Northern Mediterranean basin, Pinus halepensis Luberon Mountains in the Natural Regional Park (South of Miller (Pinaceae, Pinales) is a pioneer and expansionist France), on the basis of similar global index (climatic and species that colonizes abandoned agricultural lands char- topoedaphic conditions), by using a model developed by acterized by high biodiversity (Roche and Tatoni 1995). CEMAGREF (Ripert and Vennetier 2002). All sites Owing to its richness of secondary metabolites (Macchioni featured deep agricultural soils (>1 m) of Rendoll in “Soil et al. 2003), P. halepensis mayplayanimportantrolein Taxonomy” (Soil Survey Staff 1999) with no slope and plant succession through several processes. For example, high fertility for P. halepensis. secondary compounds (terpenoids and/or phenolic com- pounds) can affect root symbionts and site quality, by Plant Material Collection and Aqueous Leachates interfering with decomposition, mineralization, and humi- Preparation Needles and roots were collected from five fication (Kainulainen and Holopainen 2002). They can also individuals at each site. Needles were harvested from the be involved with interspecific competition phenomena entire tree crown, and roots were sampled in close proximity through allelopathic interactions (Rice 1984). Indeed, to the pines (diam <1.5 m). Just after harvest, needle and root P. halepensis may inhibit seedling establishment of extracts were soaked in water (50 g fw of tissue in 250 ml of various species in pine stands, suggesting the allelopathic distilled water). Extracts were done at room temperature (18°– nature of litter, leaf leatchates, and/or root exudates 20°C) and kept in darkness for 24 h. Needle extracts simulated (Fernandez et al. 2006, 2008;Navarro-Canoetal.2009). leaf leaching, while root extracts simulated root exudates. Other conifers such as Pinus sylvestris L. (Bulut and Bioassays. The phytotoxicity of aqueous extracts was Demir 2007), P. densiflora (Sieb. et Zucc.) (Kato-Noguchi tested in previous studies with Lactuca sativa seeds, et al. 2009), Picea abies (L.) Karst. (Pellissier 1994), or Linum strictum (allelopathy, Fernandez et al. 2006), and Picea mariana (Mill.) (Mallik and Newton 1988)also Pinus halepensis (autotoxicity, Fernandez et al. 2008). In have allelopathic potential. these studies, phytotoxicity was analyzed in terms of Allelopathic potential may be modified by several germination rate and seedling growth (roots and hypocotyles) factors such as the age of the donor plant (Inderjit and (Fernandez et al. 2006, 2008), but also in terms of sapling Asakawa 2001). Therefore, to understand the role of growth (Monnier et al. 2008). P. halepensis in secondary succession, it is essential to evaluate whether allelochemicals vary in diversity, Chemical Analyses amount, or function in different plant organs (needles vs. roots), and at different successional stages. For this reason, Instrumentation GC-MS analyses were performed on a we analyzed the composition of secondary metabolites in Hewlett-Packard 6890 GC coupled with an HP5973N Mass aqueous extracts of roots and needles of P. halepensis by Selective Detector. The GC was equipped with an HP-5MS searching for polar (fatty acids and phenolic compounds) capillary column (30 m×0.25 mm× 0.25µm—J&W Scien- and less polar compounds (terpenoids) known to be tific). Samples were injected with an ALS 7673 Automatic allelopathic (Rice 1984), and we determine whether the Injector in splitless mode (2µl for 1 min) for polar chemical diversity of aqueous extracts depends on compounds (phenolics and fatty acids), and in pulsed changes in the age of P. halepensis stands. splitless mode (5µl at 25 psi for 1 min) for less polar compounds (terpenoids). Purge flow was set to 50 ml/min after 1 min, except for qualitative studies of polar Methods and Materials compounds, for which 30 ml/min were used. Helium (99.995%) was used as carrier gas. A constant flow of Samples Sites In order to evaluate the variability and 1 ml/min was maintained throughout the runs. Three allelopathic potential of P. halepensis in relation to different oven temperatures were used for qualitative and different stages of secondary succession, three age classes quantitative studies of polar compounds, and
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