Despite plant secondary metabolites being major determinants of species interactions and ecosystem processes, their role in the maintenance of biodiversity has received little attention. In order to investigate the relationship between chemical and biological diversity in a natural ecosystem, we considered the impact of chemical diversity in individual Scots pine trees (Pinus sylvestris) on species richness of associated ground vegetation. Scots pine trees show substantial genetically determined constitutive variation between individuals in concentrations of a group of secondary metabolites, the monoterpenes. When the monoterpenes of particular trees were assessed individually, there was no relationship with species richness of associated ground flora. However, the chemical diversity of monoterpenes of individual trees was significantly positively associated with the species richness of the ground vegetation beneath each tree, mainly the result of an effect among the non-woody vascular plants. This correlation suggests that the chemical diversity of the ecosystem dominant species has an important role in shaping the biodiversity of the associated plant community. The extent and significance of this effect, and its underlying processes require further investigation.

Abstract:We identify and assess the relative importance of the principal factors influencing the release of dissolved organic carbon (DOC) and dissolved forms of nitrogen (N) from a small upland headwater dominated by podzolic soils during a sequence of autumn runoff events. We achieve this by subjecting high-resolution hydrometeorological and hydrochemical data to an R-mode principal component factor analysis and a stepwise multivariate regression analysis. We find that the release of DOC and N is influenced by four principal factors, namely event magnitude, soil water flow through the Bs horizon, the length of time since the soil profile was last flushed, and rewetting of the H horizon. The release of DOC and dissolved organic nitrogen (DON) is most strongly influenced by the combination of event magnitude and soil water flow through the Bs horizon, and to a lesser extent by the length of time since the soil profile was last flushed. Rewetting of the H horizon also influences the release of DOC, but this is not the case for DON. The release of nitrate (NO 3 -N) is most strongly influenced by the combination of the length of time since the soil profile was last flushed and rewetting of the H horizon, and to a lesser extent by event magnitude. Soil water flow through the Bs horizon does not influence the release of NO 3 -N. We argue that the mechanisms by which the above factors influence the release of DOC and N are probably strongly associated with moisture-dependent biological activity, which governs the turnover of organic matter in the soil and limits the availability of NO 3 -N in the soil for leaching. We conclude that the release of DOC and N from upland headwaters dominated by podzolic soils is largely controlled by the variable interaction of hydrometeorological factors and moisture-dependent biological processes, and that a shift in climate towards drier summers and wetter winters may result in the release of DOC and N becoming increasingly variable and more episodic in the future.

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.

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