Search

IntroductionEuropean mistletoe, Viscum album L., is a hemiparasite that can infect various tree species, yet our understanding of its physiological interactions with host species is limited.MethodsNine mistletoe–host pairs (i.e. V. album ssp. album growing on nine different broadleaf tree species) under different growth conditions in central Switzerland were selected to examine the carbon, water and nutrient relationships between mistletoe and its hosts. We measured leaf morphological traits, isotopic compositions (δ13C and δ15N), concentrations of non-structural carbohydrates (NSC) and specific compounds (i.e. mobile sugars and starch), and macronutrients (i.e. N, P, K, Ca, Mg, S) in leaf and xylem tissues of both mistletoe and its hosts.Results and DiscussionThere were only non-significant relationships between NSC concentrations in mistletoe and in its host species across the nine mistletoe–host pairs, suggesting the carbon condition of V. album ssp. album is determined by both the heterotrophic carbon transfer and self-photosynthetic capacity among different mistletoe-host pairs. However, mistletoe leaf morphological traits (single leaf area and mass, and leaf mass per unit leaf area) did not change across the nine mistletoe–host pairs, and mistletoe leaf δ13C, water content and macronutrient concentrations were linearly correlated with those in the host leaves. Macronutrients showed accumulations in mistletoe across the nine pairs. Further, tissue N concentrations were significantly higher in mistletoe grown on N-fixing hosts than on non-N-fixing hosts. Finally, leaf N:P in mistletoe was significantly correlated with the ratio in the host across the nine mistletoe–host pairs. Overall, our results indicate strong relationships between mistletoe and its hosts for water- and nutrient-related traits, but not for carbon-related traits, which demonstrates that V. album ssp. album can adjust its physiology to survive on different deciduous tree species hosts and under different site conditions.

Many carbon-related physiological questions in plants such as carbon (C) limitation or starvation have not yet been resolved thoroughly due to the lack of suitable experimental methodology. As a first step towards resolving these problems, we conducted infusion experiments with bonsai trees (Ficus microcarpa) and young maple trees (Acer pseudoplatanus) in greenhouse, and with adult Scots pine trees (Pinus sylvestris) in the field, that were “fed” with 13C-labelled glucose either through the phloem or the xylem. We then traced the 13C-signal in plant organic matter and respiration to test whether trees can take up and metabolize exogenous sugars infused. Ten weeks after infusion started, xylem but not phloem infusion significantly increased the δ13C values in both aboveground and belowground tissues of the bonsai trees in the greenhouse, whereas xylem infusion significantly increased xylem δ13C values and phloem infusion significantly increased phloem δ13C values of the adult pines in the field experiment, compared to the corresponding controls. The respiration measurement experiment with young maple trees showed significantly increased δ13C-values in shoot respired CO2 at the time of four weeks after xylem infusion started. Our results clearly indicate that trees do translocate and metabolize exogenous sugars infused, and because the phloem layer is too thin, and thus xylem infusion can be better operated than phloem infusion. This tree infusion method developed here opens up new avenues and has great potential to be used for research on the whole plant C balance and its regulation in response to environmental factors and extreme stress conditions.

It is still unknown whether the previous summer season drought and fertilization will affect the winter non-structural carbohydrate (NSC) reserves, spring leaf development, and mortality of trees in the next year. We, therefore, conducted an experiment with Quercus pubescens (downy oaks) saplings grown under four drought levels from field capacity (well-watered; ~25% volumetric water content) to wilting point (extreme drought; ~6%), in combination with two fertilizer treatments (0 vs. 50 kg/ha/year blended) for one growing season to answer this question. We measured the pre- and post-winter NSC, and calculated the over-winter NSC consumption in storage tissues (i.e. shoots and roots) following drought and fertilization treatment, and recorded the spring leaf phenology, leaf biomass, and mortality next year. The results showed that, irrespective of drought intensity, carbon reserves were abundant in storage tissues, especially in roots. Extreme drought did not significantly alter NSC levels in tissues, but delayed the spring leaf expansion and reduced the leaf biomass. Previous season fertilization promoted shoot NSC use in extreme drought-stressed saplings over winter (showing reduced carbon reserves in shoots after winter), but it also showed positive effects on survival next year. We conclude that: (1) drought-stressed downy oak saplings seem to be able to maintain sufficient mobile carbohydrates for survival, (2) fertilization can alleviate the negative effects of extreme drought on survival and recovery growth of tree saplings.

Pine mistletoe is a hemiparasitic shrub that can produce its own photosynthates. There is a lack of knowledge about the interaction of mistletoe and host under varying environmental condition that might influence carbon gain and allocation. In a 13C-pulse labeling experiment with mature Pinus sylvestris (pine) infected by mistletoes grown in naturally dry or irrigated conditions, (1) mistletoe clusters were shielded from 13CO2 added, and (2) mistletoes or host needles were removed to manipulate the local assimilate and water availability. No 13C signal was found in shielded mistletoes, indicating no carbon transfer from the host to the mistletoe. When the pine needles were removed from girdled branches, no 13C signal was found in the host tissues, implying no carbon transfer from mistletoe to the host. However, mistletoes on needle-removed pine trees accumulated more labeled assimilates and had higher non-structural carbohydrate (NSC) concentrations only under naturally dry conditions but not in irrigated plots. Our results suggest that mistletoes show full carbon autonomy, as they neither receive carbon from nor provide carbon resource to the host trees. Moreover, the high assimilation capacity of mistletoes seems to be constrained by the host water use under dry conditions, suggesting that drought stress is not only negatively impacting trees but also mistletoes. Therefore, we conclude that the hemiparasites live on their own in terms of carbon gain which, however, depends on the water provided by the host tree.

Climate-induced tree mortality became a global phenomenon during the last century and it is expected to increase in many regions in the future along with a further increase in the frequency of drought and heat events. However, tree mortality at the ecosystem level remains challenging to quantify since long-term, tree-individual, reliable observations are scarce. Here, we present a unique data set of monitoring records from 276 permanent plots located in 95 forest stands across Switzerland, which include five major European tree species (Norway spruce, Scots pine, silver fir, European beech, and sessile and common oak) and cover a time span of over one century (1898–2013), with inventory periods of 5–10 years. The long-term average annual mortality rate of the investigated forest stands was 1.5%. In general, species-specific annual mortality rates did not consistently increase over the last decades, except for Scots pine forests at lower altitudes, which exhibited a clear increase of mortality since the 1960s. Temporal trends of tree mortality varied also depending on diameter at breast height (DBH), with large trees generally experiencing an increase in mortality, while mortality of small trees tended to decrease. Normalized mortality rates were remarkably similar between species and a modest, but a consistent and steady increasing trend was apparent throughout the study period. Mixed effects models revealed that gradually changing stand parameters (stand basal area and stand age) had the strongest impact on mortality rates, modulated by climate, which had increasing importance during the last decades. Hereby, recent climatic changes had highly variable effects on tree mortality rates, depending on the species in combination with abiotic and biotic stand and site conditions. This suggests that forest species composition and species ranges may change under future climate conditions. Our data set highlights the complexity of forest dynamical processes such as long-term, gradual changes of forest structure, demography and species composition, which together with climate determine mortality rates.

Drought is a major environmental constraint to trees, causing severe stress and thus adversely affecting their functional integrity. European beech (Fagus sylvatica L.) is a key species in mesic forests that is commonly expected to suffer in a future climate with more intense and frequent droughts. Here, we assessed the seasonal response of leaf physiological characteristics of beech saplings to drought and drought release to investigate their potential to recover from the imposed stress and overcome previous limitations. Saplings were transplanted to model ecosystems and exposed to a simulated summer drought. Pre-dawn water potentials (ψpd), stomatal conductance (gS), intercellular CO2 concentration (ci), net-photosynthesis (AN), PSII chlorophyll fluorescence (PItot), non-structural carbohydrate concentrations (NSC; soluble sugars, starch) and carbon isotope signatures were measured in leaves throughout the growing season. Pre-dawn water potentials (ψpd), gS, ci, AN, and PItot decreased as drought progressed, and the concentration of soluble sugars increased at the expense of starch. Carbon isotopes in soluble sugars (δ13CS) showed a distinct increase under drought, suggesting, together with decreased ci, stomatal limitation of AN. Drought effects on ψpd, ci, and NSC disappeared shortly after re-watering, while full recovery of gS, AN, and PItot was delayed by 1 week. The fast recovery of NSC was reflected by a rapid decay of the drought signal in δ13C values, indicating a rapid turnover of assimilates and a reactivation of carbon metabolism. After recovery, the previously drought-exposed saplings showed a stimulation of AN and a trend toward elevated starch concentrations, which counteracted the previous drought limitations. Overall, our results suggest that the internal water relations of beech saplings and the physiological activity of leaves are restored rapidly after drought release. In the case of AN, stimulation after drought may partially compensate for limitations on photosynthetic activity during drought. Our observations suggest high resilience of beech to drought, contradicting the general belief that beech is particularly sensitive to environmental stressors.