A central issue in our understanding of the evolution of the diversity of plant secondary metabolites (PSMs) is whether or not compounds are functional, conferring an advantage to the plant, or non-functional. We examine the hypothesis that the diversity of monoterpene PSMs within a plant species (Scots pine Pinus sylvestris) may be explained by different compounds acting as defences against high-impact herbivores operating at different life stages. We also hypothesize that pairwise coevolution, with uncorrelated interactions, is more likely to result in greater PSM diversity, than diffuse coevolution. We tested whether up to 13 different monoterpenes in Scots pine were inhibitory to herbivory by slugs (Arion ater), bank voles (Clethrionomys glareolus), red deer (Cervus elaphus) and capercaillie (Tetrao urogallus), each of which attack trees at a different life stage. Plants containing more α-pinene were avoided by both slugs and capercaillie, which may act as reinforcing selective agents for this dominant defensive compound. Herbivory by red deer and capercaillie were, respectively, weakly negatively associated with δ3-carene, and strongly negatively correlated with the minor compound β-ocimene. Three of the four herbivores are probably contributory selective agents on some of the terpenes, and thus maintain some, but by no means all, of the phytochemical diversity in the species. The correlated defensive function of α-pinene against slugs and capercaillie is consistent with diffuse coevolutionary processes.

/pdf/do-multiple-herbivores-maintain-chemical-diversity-of-scots-46m3f3uiac.pdf

Do multiple herbivores maintain chemical diversity of Scots pine monoterpenes

Generalist parasitoids are well-known to be able to cope with the high genotypic and phenotypic plasticity of plant volatiles by learning odours during their host encounters In contrast, specialised parasitoids often respond innately to host-specific cues Previous studies have shown that females of the specialised egg parasitoid Chrysonotomyia ruforum Krausse (Hymenoptera: Eulophidae) are attracted to volatiles from Pinus sylvestris L induced by the egg deposition of its host Diprion pini L (Hymenoptera: Diprionidae), when they have previously experienced pine twigs with host eggs In this study we investigated by olfactometer bioassays how specifically C ruforum responded to oviposition-induced plant volatiles Furthermore, we studied whether parasitoids show an innate response to oviposition-induced pine volatiles Naive parasitoids were not attracted to oviposition-induced pine volatiles The attractiveness of volatiles from pines carrying eggs was shown to be specific for the pine and herbivore species, respectively (species specificity) We also tested whether not only oviposition, but also larval feeding, induces attractive volatiles (developmental stage specificity) The feeding of D pini larvae did not induce the emission of P sylvestris volatiles attractive to the egg parasitoid Our results show that a specialist egg parasitoid does not innately show a positive response to oviposition-induced plant volatiles, but needs to learn them Furthermore, the results show that C ruforum as a specialist does not learn a wide range of volatiles as some generalists do, but instead learns only a very specific oviposition-induced plant volatile pattern, ie, a pattern induced by the most preferred host species laying eggs on the most preferred food plant

Choosy egg parasitoids: specificity of oviposition-induced pine volatiles exploited by an egg parasitoid of pine sawflies

In this review we summarized recent historical records and molecular studies on evolutionary history and phylogeography of Scots pine with focus on the European highly fragmented distribution area of the species. Fossilized pollen, plant micro- and macrofossil records provided evidences on the large-scale species’ range shifts and demographic changes during the Quaternary. Populations of Scots pine were documented both in the glacial (incl. full glaciation) and interglacial periods. Recolonization of Europe after the glaciation originated from the (Sub) Mediterranean areas like the Balkan Peninsula but also from around the Eastern Alps and the surroundings of the Danube plain. Fennoscandia and northern European Baltic regions were most probably colonized from two main directions, from Western Europe and from the Russian Plain. Modern history of Scots pine was hardly affected by anthropogenic activities that started to strengthen in the Bronze and Iron Age. Along with the fossil records, molecular genetic tools were used to infer the origin and putative history including migration, differentiation and demography of the species. In this paper we compiled the major publications (30) of molecular genetic studies of the past 20 years derived from distinctly inherited organelle genomes (mitochondrial, chloroplast, nuclear) revealed by different marker systems (mtDNA-cox1, -nad1, -nad3, -nad7, ISSR, cpSSR, nSSR, B-SAP, SNP). It is important to consider that different phylogeographic patterns can be drawn by the analysis of different DNA marker types. Accordingly the use of more than one marker simultaneously outlines the most sophisticated phylogeographical pattern on the genetic lineages and can reveal high differentiation of the European distribution. Combined marker systems and markers derived from coding sequences have also been used to detect species’ phylogeographic patterns, but these were rarely applied to Scots pine. Although new molecular techniques can provide higher resolution data for populations, the reviewed results can shape the direction of further studies.

Evolutionary history and phylogeography of Scots pine (Pinus sylvestris L.) in Europe based on molecular markers

Pinus sylvestris is widely distributed in Europe and Asia, and Portugal constitutes its westernmost limit. The reduction of the Portuguese forest area of resinous species has provoked strong constraints on wood industries supply. Therefore, an increase in Scots pine area might be important, namely by reforestations of higher altitude areas, where Pinus pinaster has great difficulties to vegetate and where the risk of pinewood nematode attack is smaller. However, large gaps remain in the knowledge of Pinus sylvestris wood characteristics growing in Portugal. To address this question, the radial wood density and growth were evaluated by X-ray microdensitometric technique, sampling 100 adult trees from five representative sites of P. sylvestris distribution area in Portugal. The results revealed that Portuguese Pinus sylvestris shows good radial growth and denser wood than those found in northern European regions. Among the Portuguese stands, sites at a lower altitude (Geres and Marao) exhibited denser wood. Regarding density components, it was verified that the differences among sites were more significant in latewood, while the differences between trees/sites were most expressive in earlywood. These facts induce a higher genetic control in earlywood characteristics and a greater dependence of latewood components on environmental and climatic effects. Regarding growth components, Trees and Rings effects were more noticeable than Site effect. Concerning radial patterns, Portuguese Pinus sylvestris shows a downward trend in the first years after the pith, followed by an increase in latter rings for the density traits, while the radial variation of ring width is expressed by a tendency of decrease from the pith to the cambium. Compared to other European regions, Portuguese Pinus sylvestris reveals good wood quality features, namely higher density and ring width values. However, compared to Portuguese Pinus pinaster it shows a relatively lower density and identical or relatively lower radial growth. Scots pine could be a good solution for future reforestations of Portuguese mountainous areas, less favorable to other species.

Within and between-tree variation of wood density components in Pinus sylvestris at five sites in Portugal

Electrophoresis of Enzymes: Laboratory methods: G.M. Rothe Springer Verlag, Berlin, 1994 (ISBN 3-540-58114). 305 pp.

SUMMARY Genetic diversity is the basis for ecological biodiversity; it can only be assessed by using markers which reflect the variation present within the genome. Biochemical plant constituents interact with other components of the ecosystem and are thus of greater value as adaptive markers than as truly genotypic characters; however, the terpene components of coniferous resin systems are under strong genetic control and are valuable population diversity indicators. Molecular markers are impervious to environmental modification and therefore represent the genotype most faithfully. A survey is presented of the applications of the major biochemical and molecular marker systems in the analysis of Scots pine populations, together with indications of the advantages and limitations of each category of marker. The unusually high phenotypic diversity evident within the species at the morphological and physiological levels, both within populations and between geographical regions, is further substantiated by the degree of genotypic diversity evident from biochemical and molecular analysis. Molecular markers currently available mostly measure selectively neutral variation; among the most important needs for the future is the development of molecular markers for adaptive traits.

Genetic markers: tools for identifying and characterising scots pine populations

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Genetic markers: tools for identifying and characterising scots pine populations