Importance of tannins for responses of aspen to anthropogenic nitrogen enrichment
Abstract: Boreal forests are often strongly nitrogen (N) limited. However, human activities are leading to increased N inputs into these ecosystems, through atmospheric N deposition and forest fertilization. N input into boreal forests can promote net primary productivity, increase herbivore and pathogen damage, and shift plant species composition and community structure. Genetic diversity has been suggested as a key mechanism to promote a plant species’ stability within communities in response to environmental change. Within any plant population, specific traits (e.g. growth and defense traits) can vary substantially among individuals, and a greater variation in traits may increase chances for the persistence of at least some individuals of a population, when environmental conditions change. One aspect of plant chemistry that can greatly vary among different genotypes (GTs) are condensed tannin (CTs). These secondary metabolites have been suggested to affect plant performance in many ways, e.g. through influencing plant growth, the interactions of plants with herbivores and pathogens, and through affecting litter decomposition, and hence the return of nutrients to plants. To investigate how genotypic variation in foliar CT production may mediate the effects that anthropogenic N enrichment can have on plant performance and litter decomposition, I performed a series of experiments. For these experiments, aspen (Populus tremula) GTs with contrasting abilities to produce foliar CTs (i.e. low- vs. high-tannin producers) were grown under 3 N conditions, representing ambient N (+0 kg ha-1), upper level atmospheric N deposition (+15 kg ha-1), and forest fertilization rates (+150 kg ha-1). This general experimental set-up was once established in a field-like environment, from which natural enemies were excluded, and once in a field, in which enemies were present. In my first two studies, I investigated tissue chemistry and plant performance in both environments. I observed that foliar CT levels decreased in response to N in the enemy‑free environment (study I), but increased with added N when enemies were present (study II). These opposing responses to N may be explained by differences in soil N availability in the two environments, or by induction of CTs after enemy attack. Enemy damage generally increased in response to N, and was higher in low-tannin than in high-tannin plants across all N levels. Plant growth of high‑tannin plants was restricted under ambient and low N conditions, probably due to a trade-off between growth and defense. This growth constraint for high‑tannin plants was weakened, when high amounts of N were added (study I and II), and when enemy levels were sufficiently high, so that benefits gained through defense could outweigh the costs of defense production (study II). Despite those general responses of low- and high‑tannin producers to added N, I also observed a number of individual responses of GTs to N addition, which in some case were not connected to the intrinsic ability of the GTs to produce foliar CTs. In study III, gene expression levels in young leaves and phenolic pools of the plants that were grown in the enemy‑free environment were studied. This study revealed that gene control over the regulation of the phenylpropanoid pathway (PPP) was distributed across the entire pathway. Moreover, PPP gene expression was higher in high-tannin GTs than in low‑tannin GTs, particularly under ambient N. At the low N level, gene expressions declined for both low- and high-tannin producers, whereas at the high N level expression at the beginning and the end of the PPP was upregulated and difference between tannin groups disappeared. Furthermore, this study showed that phenolic pools were frequently uncorrelated, and that phenolic pools were only to some extent related to tannin production and gene expression. In study IV, I investigated the decomposability of litter from the field plants. I found that N enrichment generally decreased mass loss, but there was substantial genetic variation in decomposition rates, and GTs were differentially responsive to added N. Study IV further showed that CTs only had a weak effect on decomposition, and other traits, such as specific leaf area and the lignin:N ratio, could better explain genotypic difference in mass loss. Furthermore, N addition caused a shift in which traits most strongly influenced decomposition rates. Collectively, the result of these studies highlight the importance of genetic diversity to promote the stability of species in environments that experience anthropogenic change.
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