Microbial processing of plant remains is co-limited by multiple nutrients in global grasslands.

Microbial processing of aggregate-unprotected organic matter inputs is key for soil fertility, long-term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro- and micro-nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak-season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains.

Physico-chemical properties of plant cuticles and their functional and ecological significance.

Most aerial plant surfaces are covered with a lipid-rich cuticle, which is a barrier for the bidirectional transport of substances between the plant and the surrounding environment. This review article provides an overview of the significance of the leaf cuticle as a barrier for the deposition and absorption of water and electrolytes. After providing insights into the physico-chemical properties of plant surfaces, the mechanisms of foliar absorption are revised with special emphasis on solutes. Due to the limited information and relative importance of the leaf cuticle of herbaceous and deciduous cultivated plants, an overview of the studies developed with Alpine conifers and treeline species is provided. The significance of foliar water uptake as a phenomenon of ecophysiological relevance in many areas of the world is also highlighted. Given the observed variability in structure and composition among, for example, plant species and organs, it is concluded that it is currently not possible to establish general permeability and wettability models that are valid for predicting liquid-surface interactions and the subsequent transport of water and electrolytes across plant surfaces.

Leaf wettability, anatomy and ultra-structure of Nothofagus antarctica and N. betuloides grown under a CO2 enriched atmosphere.

Increasing CO2 air concentration may affect wettability, anatomy and ultra-structure of leaves of Patagonian forest species, evergreen and deciduous plants potentially responding differently to such CO2 increases. In this study, we analysed the wettability, anatomy and ultra-structure of leaves of Nothofagus antarctica (deciduous) and N. betuloides (evergreen) grown under high CO2 concentrations. Leaf wettability was affected by increasing CO2, in different directions depending on species and leaf side. In both species, soluble cuticular lipid concentrations per unit leaf area raised with higher CO2 levels. Stomatal parameters (density, size of guard cells and pores) showed different responses to CO2 increasing depending on the species examined. In both species, leaf tissues showed a general trend to diminish with higher CO2 concentration. Cuticle thickness was modified with higher CO2 concentration in N. betuloides, but not in N. antarctica leaves. In both species, chloroplasts were often damaged with the increase in CO2 concentration. Our results show that several surface and internal leaf parameters can be modified in association with an increase in atmospheric CO2 concentration which may very among plant species.

Foliar Application of Potassium Salts to Olive, with Focus on Accompanying Anions.

Potassium (K) is an essential element, which is often supplied to horticultural crops via foliar spraying. Some studies have investigated the effect of different foliar-applied K compounds; however, most studies have focussed on crop quality and yield parameters, or were performed with isolated leaf cuticles. The aim of this study was to evaluate the rates of the foliar ion penetration and leaf surface deposition of 130 mM K sprays of compounds with markedly different point of deliquescence (POD) and efflorescence (POE) values, the rates having been previously estimated in climate chamber trials. Shoots of field-grown, commercial olive trees were sprayed with K-nitrate (KNO3), K-sulphate (K2SO4), K-chloride (KCl), K-phosphate (K3PO4), K-carbonate (K2CO3) and K-bicarbonate (KHCO3), and leaf samples were collected after 3 and 24 h. Cation and anion concentrations were determined in the leaf tissues, and in a preliminary leaf water wash for estimating surface-deposited ion concentrations. No significant leaf tissue K increments were recorded between the K sprays. Olive tissue anion concentrations showed different patterns, and a chloride (Cl-) increase was detected 3 h after the foliar KCl supply. On the other hand, the foliar K applications led to leaf nitrate changes regardless of the K source supplied. High amounts of K and accompanying ions were recovered in the washing liquid of the foliar K-supplied leaves. Some foliar K treatments increased the leaf surface concentration of sulphate and chloride, suggesting a potential effect on leaf cell anion extrusion. Hence, despite no evidence of foliar K uptake, an effect of leaf anion concentrations was observed, indicating that foliar nutrient sprays may influence leaf and leaf surface anion balance.

Herbivory and nutrients shape grassland soil seed banks.

Anthropogenic nutrient enrichment and shifts in herbivory can lead to dramatic changes in the composition and diversity of aboveground plant communities. In turn, this can alter seed banks in the soil, which are cryptic reservoirs of plant diversity. Here, we use data from seven Nutrient Network grassland sites on four continents, encompassing a range of climatic and environmental conditions, to test the joint effects of fertilization and aboveground mammalian herbivory on seed banks and on the similarity between aboveground plant communities and seed banks. We find that fertilization decreases plant species richness and diversity in seed banks, and homogenizes composition between aboveground and seed bank communities. Fertilization increases seed bank abundance especially in the presence of herbivores, while this effect is smaller in the absence of herbivores. Our findings highlight that nutrient enrichment can weaken a diversity maintaining mechanism in grasslands, and that herbivory needs to be considered when assessing nutrient enrichment effects on seed bank abundance.

Effect of irradiation and canopy position on anatomical and physiological features of Fagus sylvatica and Quercus petraea leaves.

Growing conditions at different tree canopy positions may significantly vary and lead to foliar changes even within the same tree. An assessment of foliar anatomy, including also epidermal features, can help us understand how plants respond to environmental factors. Working with two model tree species (i.e., Quercus petraea and Fagus sylvatica) grown at their southernmost European distribution area in Central Spain, the influence of irradiation and canopy height was examined by sampling lower canopy leaves and comparing them with fully irradiated, top canopy leaves and shaded top canopy leaves grown for months within a bag made of shade netting fabric before they sprouted. At the end of the summer, samples were collected, and several parameters were analysed. The results indicate that SLA (specific leaf area) differences are significant both between species and groups. Leaf and cuticle thickness differed significantly between groups while stomatal densities only between species. Regarding mineral concentrations, differences between species were significant for K, Mn, N and N: P ratios. It is concluded that leaf responses to environmental conditions may be variable both within the same tree and between species.

Essential Nutrient and Trace Element Foliar Resorption of Two Co-Existing Nothofagus Species Grown Under Different Environmental Conditions in Southern Patagonia.

Nutrient resorption is crucial for mineral element conservation and efficiency of forest species, but knowledge on its significance and the mechanisms involved is still limited for most species and habitats. Focusing on the harsh conditions for plant growth and survival of southern Patagonia, a field study for comparing the rate of foliar resorption of macro-, micro-nutrients, and trace elements in coexisting Nothofagus pumilio and Nothofagus antarctica forests was performed. Forests located in three contrasting productivity sites (with different soil and climatic conditions) were selected, and mature, functional versus senescent leaves of both species were collected at two different dates of the growing season. Macro- (N, P, Ca, K, S, and Mg), micronutrients (B, Cu, Fe, Mn, Zn, and Ni), and trace elements (Al, Li, Pb, Rb, Sr, Ti, and Tl) were determined in foliar tissues. The mineral element concentrations of mature and senescent leaves were used for calculating the nutrient resorption efficiency (NuR). In general, and making an average of all sites and species, macro-nutrient resorption showed a decreasing trend for N > S = K > P > Mg, being Ca the only macro-nutrient with negative values (i.e., no resorption). Resorption of the majority of the elements did not vary between species in any of the evaluated sites. Variation across sites in nutrient resorption efficiency for most macronutrients, some micronutrients, and trace elements was observed for N. antarctica, whereas N. pumilio had a similar NuR for all experimental sites. On the other hand, regardless of the site or the species, some elements were not resorbed (e.g., B, Cu, Fe, Mn, Al, and Ti). It is concluded that both Nothofagus species performed similarly concerning their nutrient conservation strategy, when coexisting in the same mixed forest. However, no evidence was gained for an increased rate of foliar NuR in association with the sites subjected to more limiting soil and climatic conditions for plant growth.

Surface Properties and Permeability to Calcium Chloride of Fagus sylvatica and Quercus petraea Leaves of Different Canopy Heights.

Plant surfaces have a considerable degree of chemical and physical variability also in relation to different environmental conditions, organs and state of development. The potential changes on plant surface properties in association with environmental variations have been little explored so far. Using two model tree species (i.e., Quercus petraea, sessile oak and Fagus sylvatica, beech) growing in 'Montejo de la Sierra Forest,' we examined various traits of the abaxial and adaxial surface of leaves of both species collected at a height of approximately 15 m (top canopy), versus 3.5-5.5 m for beech and sessile oak, lower canopy leaves. Leaf surface ultra-structure was analyzed by scanning and transmission electron microscopy, and the surface free energy and related parameter were estimated after measuring drops of 3 liquids with different degrees of polarity and apolarity. The permeability of the adaxial and abaxial surface of top and bottom canopy leaves to CaCl2 was estimated by depositing 2 drops of 3-4 µl per cm2 and comparing the concentration of Ca in leaf tissues 24 h after treatment, and also Ca and Cl concentrations in the washing liquid. Higher Ca concentrations were recorded after the application of CaCl2 drops onto the veins and adaxial blade of top canopy beech leaves, while no significant evidence for foliar Ca absorption was gained with sessile oak leaves. Surprisingly, high amounts of Cl were recovered after washing untreated, top canopy beach and sessile oak leaves with deionised water, a phenomenon which was not traced to occur on lower canopy leaves of both species. It is concluded that the surface of the two species analyzed is heterogeneous in nature and may have areas favoring the absorption of water and solutes as observed for the veins of beech leaves.

Intra-population variability in the drought response of a beech (Fagus sylvatica L.) population in the southwest of Europe.

Phenotypic variability within forest species populations is considered of special relevance for local adaptation under new environments, albeit it has been analyzed to a lesser extent than inter-population phenotypic variability. A common garden study was carried out to assess phenotypic variability in response to water stress in half-sibling families from a marginal population of Fagus sylvatica L. at its south-western range edge distribution in Europe. Two irrigation regimes were applied, well-watered (WW) seedlings and those submitted to weekly cycles of drying-rewatering of growth media. Seedling growth and their leaf functional traits were recorded during the last cycle of water stress. Most of the phenotypic changes were explained by phenotypic plasticity in response to water stress, but there was also a significant effect of family in the expression of some of the studied traits. The relationship of carbon isotope fractioning with gas exchange traits across families under WW conditions did not follow the same pattern as the phenotypic trends. The leaf net photosynthesis across families was modified by the nitrogen content on a leaf mass basis that was in turn correlated positively with leaf nitrogen isotope fractionation. The results point to an important role of leaf nitrogen in determining the intrinsic water-use efficiency (WUE) across families. Variation in WUE was ruled mainly by control of stomatal conductance to water vapor under water stress, but by leaf net photosynthesis under wet conditions. Relatively high inter-family phenotypic variability in growth and functional traits were observed. Within-population phenotypic variability, and the plasticity of some of the studied traits, is of fundamental importance to cope with the harsher environments beech will have to endure in the future at different points in its distribution range.