Plant community composition affects the species biogeochemical niche

Nutrients are essential for plant development, and their availability and stoichiometric ratios can influence the composition of plant communities. We investigated the possibility of the reverse influence: whether the conditions of contrasting species coexistence determine foliar element concentrations and plant stoichiometry, that is, species biogeochemical niche (BN). The experiment was conducted at the Ecological-Botanical Garden of the University of Bayreuth, Germany. We analyzed foliar element concentrations of two dwarf shrubs (Calluna vulgaris and Vaccinium myrtillus) and two grasses (Holcus lanatus and Arrhenatherum elatius) growing in different community compositions (monocultures and various mixed stands). Foliar nutrient concentrations and stoichiometry (taken as a proxy of species BN) were species specific; each species showed its own BN in all communities. Furthermore, V. myrtillus and H. lanatus species shifted their BN in response to changes in their community, accomplishing the "biogeochemical niche displacement" hypothesis. We conclude that plants can readjust their foliar element concentration if they grow in communities with contrasting plant composition, suggesting a differential use of element resources when the patterns of species coexistence change. These results also support the complementary niche hypothesis

Local adaptations to frost in marginal and central populations of the dominant forest tree Fagus sylvatica L. as affected by temperature and extreme drought in common garden experiments

Local adaptations to environmental conditions are of high ecological importance as they determine distribution ranges and likely affect species responses to climate change. Increased environmental stress (warming, extreme drought) due to climate change in combination with decreased genetic mixing due to isolation may lead to stronger local adaptations of geographically marginal than central populations. We experimentally observed local adaptations of three marginal and four central populations of Fagus sylvaticaL., the dominant native forest tree, to frost over winter and in spring (late frost). We determined frost hardiness of buds and roots by the relative electrolyte leakage in two common garden experiments. The experiment at the cold site included a continuous warming treatment; the experiment at the warm site included a preceding summer drought manipulation. In both experiments, we found evidence for local adaptation to frost, with stronger signs of local adaptation in marginal populations. Winter frost killed many of the potted individuals at the cold site, with higher survival in the warming treatment and in those populations originating from colder environments. However, we found no difference in winter frost tolerance of buds among populations, implying that bud survival was not the main cue for mortality. Bud late frost tolerance in April differed between populations at the warm site, mainly because of phenological differences in bud break. Increased spring frost tolerance of plants which had experienced drought stress in the preceding summer could also be explained by shifts in phenology. Stronger local adaptations to climate in geographically marginal than central populations imply the potential for adaptation to climate at range edges. In times of climate change, however, it needs to be tested whether locally adapted populations at range margins can successfully adapt further to changing conditions

Opposite metabolic responses of shoots and roots to drought

Shoots and roots are autotrophic and heterotrophic organs of plants with different physiological functions. Do they have different metabolomes? Do their metabolisms respond differently to environmental changes such as drought? We used metabolomics and elemental analyses to answer these questions. First, we show that shoots and roots have different metabolomes and nutrient and elemental stoichiometries. Second, we show that the shoot metabolome is much more variable among species and seasons than is the root metabolome. Third, we show that the metabolic response of shoots to drought contrasts with that of roots; shoots decrease their growth metabolism (lower concentrations of sugars, amino acids, nucleosides, N, P, and K), and roots increase it in a mirrored response. Shoots are metabolically deactivated during drought to reduce the consumption of water and nutrients, whereas roots are metabolically activated to enhance the uptake of water and nutrients, together buffering the effects of drought, at least at the short term

Shifts in the elemental composition of plants during a very severe drought

Diverse plant functions (e.g., growth, storage, defense and anti-stress mechanisms) use elements disproportionally. We hypothesized that plants growing under different abiotic and biotic conditions would shift their elemental compositions in response to a very severe drought. We tested this hypothesis by investigating the changes in foliar stoichiometry and species composition from a very severe drought. We also tested the effects of previous droughts (acclimation) on this response. Different species growing in the same community responded more similarly to a very severe drought than did individual species growing in different communities. The stoichiometric shifts were thus more community-dependent than species-dependent. The results also suggested that plants grown in monoculture were less stoichiometrically plastic during the drought than plants grown in a more diverse community. Previous exposure to long-term drought treatments in the same communities did not significantly affect the stoichiometric shifts during the new drought. Differential use of resources may have been responsible for these responses. Monocultured plants, which used the same resources in similar proportions, had more difficulty avoiding direct competition when the resources became scarcer. Moreover, each species tested had a particular elemental composition in all communities and climatic treatments. The differences in foliar elemental compositions were largest between plant functional groups (shrubs and grasses) and smallest among species within the same functional group. Global principal components analyses (PCAs) identified a general tendency for all species, independently of the community in which they grew, toward lower concentrations of K, N, P, Mg and S, and to higher concentrations of C and Fe as the drought advanced. This study has demonstrated the utility of analyses of differences and shifts in plant elemental composition for understanding the processes underlying the responses of plants to changes in biotic and abiotic environmental conditions

Winter warming is ecologically more relevant than summer warming in a cool-temperate grassland

Climate change affects all seasons, but warming is more pronounced in winter than summer at mid- and high latitudes. Winter warming can have profound ecological effects, which are rarely compared to the effects of summer warming, and causal explanations are not well established. We compared mild aboveground infrared warming in winter to warming in summer in a semi-natural, cool-temperate grassland in Germany for four years. Aboveground plant biomass increased following winter warming (+18%) and was unaffected by summer warming. Winter warming affected the composition of the plant community more than summer warming, favoring productive species. Winter warming increased soil respiration more than summer warming. Prolonged growing seasons and changes in plant-community composition accounted for the increased aboveground biomass production. Winter warming stimulated ecological processes, despite causing frost damage to plant roots and microorganisms during an extremely cold period when warming reduced the thermal insulation provided by snow. Future warming beyond such intermittent frosts may therefore further increase the accelerating effects of winter warming on ecological processe

Climatically controlled reproduction drives interannual growth variability in a temperate tree species

Climatically controlled allocation to reproduction is a key mechanism by which climate influences tree growth and may explain lagged correlations between climate and growth. We used continent‐wide datasets of tree‐ring chronologies and annual reproductive effort in Fagus sylvatica from 1901 to 2015 to characterise relationships between climate, reproduction and growth. Results highlight that variable allocation to reproduction is a key factor for growth in this species, and that high reproductive effort (‘mast years’) is associated with stem growth reduction. Additionally, high reproductive effort is associated with previous summer temperature, creating lagged climate effects on growth. Consequently, understanding growth variability in forest ecosystems requires the incorporation of reproduction, which can be highly variable. Our results suggest that future response of growth dynamics to climate change in this species will be strongly influenced by the response of reproduction.Additional co-authors: Ernst van der Maaten, Marieke van der Maaten‐Theunissen, Lena Muffler, Renzo Motta, Catalin‐Constantin Roibu, Ionel Popa, Tobias Scharnweber, Robert Weigel, Martin Wilmking, Christian S Zan

Biotic homogenization destabilizes ecosystem functioning by decreasing spatial asynchrony

Our planet is facing significant changes of biodiversity across spatial scales. Although the negative effects of local biodiversity (α diversity) loss on ecosystem stability are well documented, the consequences of biodiversity changes at larger spatial scales, in particular biotic homogenization, that is, reduced species turnover across space (β diversity), remain poorly known. Using data from 39 grassland biodiversity experiments, we examine the effects of β diversity on the stability of simulated landscapes while controlling for potentially confounding biotic and abiotic factors. Our results show that higher β diversity generates more asynchronous dynamics among local communities and thereby contributes to the stability of ecosystem productivity at larger spatial scales. We further quantify the relative contributions of α and β diversity to ecosystem stability and find a relatively stronger effect of α diversity, possibly due to the limited spatial scale of our experiments. The stabilizing effects of both α and β diversity lead to a positive diversity–stability relationship at the landscape scale. Our findings demonstrate the destabilizing effect of biotic homogenization and suggest that biodiversity should be conserved at multiple spatial scales to maintain the stability of ecosystem functions and services

Increased Soil Frost Versus Summer Drought as Drivers of Plant Biomass Responses To Reduced Precipitation: Results from A Globally-Coordinated Field Experiment

Reduced precipitation treatments often are used in field experiments to explore the effects of drought on plant productivity and species composition. However, in seasonally snow-covered regions reduced precipitation also reduces snow cover, which can increase soil frost depth, decrease minimum soil temperatures and increase soil freeze-thaw cycles. Therefore, in addition to the effects of reduced precipitation on plants via drought, freezing damage to overwintering plant tissues at or below the soil surface could further affect plant productivity and relative species abundances during the growing season. We examined the effects of both reduced rainfall (via rain-out shelters) and reduced snow cover (via snow removal) at 13 sites globally (primarily grasslands) within the framework of the International Drought Experiment, a coordinated distributed experiment. Plant cover was estimated at the species level and aboveground biomass was quantified at the functional group level. Among sites, we observed a negative correlation between the snow removal effect on minimum soil temperature and plant biomass production the next growing season. Three sites exhibited significant rain-out shelter effects on plant productivity, but there was no correlation among sites between the rain-out shelter effect on minimum soil moisture and plant biomass. There was no interaction between snow removal and rain-out shelters for plant biomass, although these two factors only exhibited significant effects simultaneously for a single site. Overall, our results reveal that reduced snowfall, when it decreases minimum soil temperatures, can be an important component of the total effect of reduced precipitation on plant productivity

Warming differentially influences the effects of drought on stoichiometry and metabolomics in shoots and roots

Plants in natural environments are increasingly being subjected to a combination of abiotic stresses, such as drought and warming, in many regions. The effects of each stress and the combination of stresses on the functioning of shoots and roots have been studied extensively, but little is known about the simultaneous metabolome responses of the different organs of the plant to different stresses acting at once. - We studied the shift in metabolism and elemental composition of shoots and roots of two perennial grasses, Holcus lanatus and Alopecurus pratensis, in response to simultaneous drought and warming. -These species responded differently to individual and simultaneous stresses. These responses were even opposite in roots and shoots. In plants exposed to simultaneous drought and warming, terpenes, catechin and indole acetic acid accumulated in shoots, whereas amino acids, quinic acid, nitrogenous bases, the osmoprotectants choline and glycine betaine, and elements involved in growth (nitrogen, phosphorus and potassium) accumulated in roots. Under drought, warming further increased the allocation of primary metabolic activity to roots and changed the composition of secondary metabolites in shoots. -These results highlight the plasticity of plant metabolomes and stoichiometry, and the different complementary responses of shoots and roots to complex environmental conditions