Publications (5)17.78 Total impact
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Article: Assessment and Implications of Intraspecific and Phenological Variability in Monoterpenes of Scots Pine (Pinus sylvestris) Foliage
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ABSTRACT: Scots pine populations contain individuals with widely differing amounts and composition of monoterpenes and exist as one of two chemotypes: with or without Δ3-carene. We investigated the significance for ecological studies of two types of variation in monoterpenes: (1) the inherent variability in the concentration of monoterpenes or their relative amounts in needles of seedlings, saplings, and mature trees; and (2) phenological variation in developing needles. The relative composition of needle monoterpenes in 5-year-old saplings changed during the needle development period until the final composition was reached upon needle maturity. Changes in composition depended on chemotype. Needles of the “no-Δ3-carene” chemotype had higher absolute concentrations of α-pinene, β-pinene, camphene, and total monoterpenes than “Δ3-carene” chemotype. For the “Δ3-carene” chemotype, the relative concentration of Δ3-carene during the needle growing season and immediately after emergence of seedlings was higher compared to that reached at needle maturity. Repeated removal of single needles (at weekly intervals during growth) from 5-year-old saplings did not influence the composition of monoterpenes. Within a natural Scots pine dominated woodland, 18% of mature Scots pines (N = 574) belonged to the “no-Δ3-carene” chemotype. Chemotypic variation within populations means that the statistical power with which differences in monoterpene concentrations can be detected is lower when sampling from the whole population compared to sampling within chemotypes. Reduction of this background variation and accounting for chiral variation if present, would significantly aid efficiency, interpretation, and understanding of processes in chemical and ecological research. One method for achieving this is the screening of plants for chemotypes before the establishment of experiments or field sampling regimes. We present a summary of suitable analytical methods for needle tissue that facilitates this prior screening.Journal of Chemical Ecology 04/2012; 33(3):477-491. · 2.66 Impact Factor -
Article: Do multiple herbivores maintain chemical diversity of Scots pine monoterpenes?
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ABSTRACT: 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 δ(3)-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.Philosophical Transactions of The Royal Society B Biological Sciences 05/2011; 366(1569):1337-45. · 6.40 Impact Factor -
Article: The role of genetic and chemical variation of Pinus sylvestris seedlings in influencing slug herbivory.
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ABSTRACT: This study investigated the genetic and chemical basis of resistance of Pinus sylvestris seedlings to herbivory by a generalist mollusc, Arion ater. Using feeding trials with captive animals, we examined selective herbivory by A. ater of young P. sylvestris seedlings of different genotypes and correlated preferences with seedling monoterpene levels. We also investigated the feeding responses of A. ater to artificial diets laced with two monoterpenes, Delta(3)-carene and alpha-pinene. Logistic regression indicated that two factors were the best predictors of whether seedlings in the trial would be consumed. Individual slug variation (replicates) was the most significant factor in the model; however, alpha-pinene concentration (also representing beta-pinene, Delta(3)-carene and total monoterpenes due to multicollinearity) of needles was also a significant factor. While A. ater did not select seedlings on the basis of family, seedlings not eaten were significantly higher in levels of alpha-pinene compared to seedlings that were consumed. We also demonstrated significant genetic variation in alpha-pinene concentration of seedlings between different families of P. sylvestris. Nitrogen and three morphological seedling characteristics (stem length, needle length and stem diameter) also showed significant genetic variation between P. sylvestris families. Artificial diets laced with high (5 mg g(-1) dry matter) quantities of either Delta(3)-carene or alpha-pinene, were eaten significantly less than control diets with no added monoterpenes, supporting the results of the seedling feeding trial. This study demonstrates that A. ater selectively feed on P. sylvestris seedlings and that this selection is based, in part, on the monoterpene concentration of seedlings. These results, coupled with significant genetic variation in alpha-pinene concentration of seedlings and evidence that slug herbivory is detrimental to P. sylvestris fitness, are discussed as possible evidence for A. ater as a selective force on the evolution of defensive chemistry in P. sylvestris.Oecologia 06/2007; 152(1):82-91. · 3.41 Impact Factor -
Article: Assessment and implications of intraspecific and phenological variability in monoterpenes of Scots pine (Pinus sylvestris) foliage.
[show abstract] [hide abstract]
ABSTRACT: Scots pine populations contain individuals with widely differing amounts and composition of monoterpenes and exist as one of two chemotypes: with or without delta3-carene. We investigated the significance for ecological studies of two types of variation in monoterpenes: (1) the inherent variability in the concentration of monoterpenes or their relative amounts in needles of seedlings, saplings, and mature trees; and (2) phenological variation in developing needles. The relative composition of needle monoterpenes in 5-year-old saplings changed during the needle development period until the final composition was reached upon needle maturity. Changes in composition depended on chemotype. Needles of the "no-delta3-carene" chemotype had higher absolute concentrations of alpha-pinene, beta-pinene, camphene, and total monoterpenes than "delta3-carene" chemotype. For the "delta3-carene" chemotype, the relative concentration of delta3-carene during the needle growing season and immediately after emergence of seedlings was higher compared to that reached at needle maturity. Repeated removal of single needles (at weekly intervals during growth) from 5-year-old saplings did not influence the composition of monoterpenes. Within a natural Scots pine dominated woodland, 18% of mature Scots pines (N=574) belonged to the "no-delta3-carene" chemotype. Chemotypic variation within populations means that the statistical power with which differences in monoterpene concentrations can be detected is lower when sampling from the whole population compared to sampling within chemotypes. Reduction of this background variation and accounting for chiral variation if present, would significantly aid efficiency, interpretation, and understanding of processes in chemical and ecological research. One method for achieving this is the screening of plants for chemotypes before the establishment of experiments or field sampling regimes. We present a summary of suitable analytical methods for needle tissue that facilitates this prior screening.Journal of Chemical Ecology 04/2007; 33(3):477-91. · 2.66 Impact Factor -
Article: Behavioral strategies of mammal herbivores against plant secondary metabolites: the avoidance-tolerance continuum.
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ABSTRACT: We review the evidence for behavioral avoidance of plant secondary metabolites (PSMs) and identify how, and the circumstances under which it occurs. Behavioral strategies of avoidance of PSM can only be fully understood in relation to the underlying physiological processes or constraints. There is considerable evidence that animals learn to avoid PSMs on the basis of negative postingestive effects. The extent to which this process determines foraging choices is limited by the ability of animals to experience the consequences of their behaviors and associate particular cues in foods with their specific effects in the body. The proposed learning mechanisms require that animals must at least "sample" plants that contain PSMs. They do not completely avoid PSMs, but there is evidence that they restrict their ingestion to within limits that they are physiologically able to tolerate, and that the amounts of PSM ingested result from a balance between toxicological considerations and the nutrient content of the plant material. These limits are influenced by the kinetics of PSM elimination, which underlies patterns of bite and patch selection from plant parts to landscapes. We suggest that altering spatial location of feeding (to alternative food patches or alternative foods within patches, including plant parts), and temporal distribution of feeding activity, by either cessation of feeding or by continuing to feed, but on alternative foods, can both lead to reduction of the intake and toxic effects of PSMs. We propose that the strategy of avoidance or reduction of intake of PSMs coevolved with the animal's ability to physiologically tolerate their ingestion, and that avoidance and tolerance are inversely related (the avoidance-tolerance continuum). The animals' propensity and ability to seek alternatives also vary with the dispersion of their food resources. Further work is required to test these proposals and integrate temporal and spatial aspects of foraging behavior and its nutritional consequences in relation to PSMs.Journal of Chemical Ecology 07/2006; 32(6):1115-32. · 2.66 Impact Factor
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2007
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Bangor University
Bangor, WLS, United Kingdom
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