Global patterns of leaf construction traits and their covariation along climate and soil environmental gradients.

Leaf functional traits and their covariation underlie plant ecological adaptations along environmental gradients, but there is limited information on the global covariation patterns of key leaf construction traits. To explore how leaf construction traits co-vary across diverse climate and soil environmental conditions, we compiled a global dataset including cell wall mass per unit leaf mass (CWmass ), leaf carbon (C) and calcium (Ca) concentrations, and specific leaf area (SLA) for 2348 angiosperm species from 340 sites world-wide. Our results demonstrated negative correlations between leaf C and Ca concentrations and between leaf C and SLA across diverse nongraminoid angiosperms. Leaf C concentration increased with increasing mean annual temperature (MAT) and mean annual precipitation (MAP) and with decreasing soil pH and calcium carbonate (CaCO3 ) concentration, whereas leaf Ca concentration and SLA exhibited the opposite responses to these environmental variables. The covariations of leaf Ca-C and of leaf SLA-C were stronger in habitats with lower MAT and MAP, and/or higher soil CaCO3 content. This global-scale analysis demonstrates that the leaf C and Ca concentrations and SLA together govern the C and biomass investment strategies in leaves of nongraminoids. We conclude that environmental conditions strongly shape leaf construction traits and their covariation patterns.

The Response of Plants and Mycorrhizal Fungi to Nutritionally-Heterogeneous Environments Are Regulated by Nutrient Types and Plant Functional Groups.

Nutrient type and plant functional group are both important in influencing proliferation of roots or hyphae and their benefit to plant growth in nutritionally heterogeneous environments. However, the studies quantifying relative importance of roots vs. hyphae affecting the plant response to nutrient heterogeneity are lacking. Here, we used meta-analysis based on 879 observations from 66 published studies to evaluate response patterns of seven variables related to growth and morphological traits of plants and mycorrhizal fungi in nutritionally heterogeneous environments. We found that phosphorus [P] and organic fertilizer [OF] supply significantly increased shoot (+18.1 and +25.9%, respectively) and root biomass (+31.1 and +23.0%, respectively) and root foraging precision (+11.8 and +20.4%, respectively). However, there was no significant difference among functional groups of herbs (grasses, forbs, and legumes), between herbs and woody species, and between arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) tree species in the shoot, root and mycorrhizal fungi responses to nutrient heterogeneity, except for root biomass and root foraging precision among grasses, forbs, and legumes, and mycorrhizal hyphal foraging precision between AM and ECM tree species. Root diameter was uncorrelated with neither root foraging precision nor mycorrhizal hyphal foraging precision, regardless of mycorrhizal type or nutrient type. These results suggest that plant growth and foraging strategies are mainly influenced by nutrient type, among other factors including plant functional type and mycorrhizal type.

Relationships Between Leaf Carbon and Macronutrients Across Woody Species and Forest Ecosystems Highlight How Carbon Is Allocated to Leaf Structural Function.

Stoichiometry of leaf macronutrients can provide insight into the tradeoffs between leaf structural and metabolic investments. Structural carbon (C) in cell walls is contained in lignin and polysaccharides (cellulose, hemicellulose, and pectins). Much of leaf calcium (Ca) and a fraction of magnesium (Mg) were further bounded with cell wall pectins. The macronutrients phosphorus (P), potassium (K), and nitrogen (N) are primarily involved in cell metabolic functions. There is limited information on the functional interrelations among leaf C and macronutrients, and the functional dimensions characterizing the leaf structural and metabolic tradeoffs are not widely appreciated. We investigated the relationships between leaf C and macronutrient (N, P, K, Ca, Mg) concentrations in two widespread broad-leaved deciduous woody species Quercus wutaishanica (90 individuals) and Betula platyphylla (47 individuals), and further tested the generality of the observed relationships in 222 woody eudicots from 15 forest ecosystems. In a subsample of 20 broad-leaved species, we also analyzed the relationships among C, Ca, lignin, and pectin concentrations in leaf cell walls. We found a significant leaf C-Ca tradeoff operating within and across species and across ecosystems. This basic relationship was explained by variations in the share of cell wall lignin and pectin investments at the cell scale. The C-Ca tradeoffs were mainly driven by soil pH and mean annual temperature and precipitation, suggesting that leaves were more economically built with less C and more Ca as soil pH increased and at lower temperature and lower precipitation. However, we did not detect consistent patterns among C-N, and C-Mg at different levels of biological organization, suggesting substantial plasticity in N and Mg distribution among cell organelles and cell protoplast and cell wall. We observed two major axes of macronutrient differentiation: the cell-wall structural axis consisting of protein-free C and Ca and the protoplasm metabolic axis consisting of P and K, underscoring the decoupling of structural and metabolic elements inherently linked with cell wall from protoplasm investment strategies. We conclude that the tradeoffs between leaf C and Ca highlight how carbon is allocated to leaf structural function and suggest that this might indicate biogeochemical niche differentiation of species.

The persistence of bacterial diversity and ecosystem multifunctionality along a disturbance intensity gradient in karst soil.

Extensive, progressive rock emergence causes localized variations in soil biogeochemical and microbial properties that may influence the capacity for the regeneration of degraded karst ecosystems. It is likely that karst ecosystem recovery relies on the persistence of soil functions at the microbial scale, and we aimed to explored the role of interactions between soil bacterial taxa and identify keystone species that deliver key biogeochemical functions, i.e. carbon (C) and nutrient (nitrogen, N and phosphorus, P) cycling. We applied high-throughput sequencing and phylogenetic molecular ecological network approaches to topsoils sampled at rock-soil interfaces and adjacent bulk soil along an established gradient of land-use intensity in the Chinese Karst Critical Zone Observatory. Bacterial α-diversity was greater under increased perturbation and at the rock-soil interface compared to bulk soils under intensive cultivation. However, bacterial ecological networks were less intricate and connected fewer keystone taxa as human disturbance increased and at the rock-soil interface. Co-occurrence within the bacterial community in natural primary forest soils was 13% larger than cultivated soils. The relative abundances of keystone taxa Acidobacteria, Bacteroidetes and Chloroflexi increased with land-use intensity, while Proteobacteria, Actinobacteria and Verrucomicrobia decreased by up to 6%. In general, Bacteroidetes, Verrucomicrobia and Chlorobi were related to C-cycling, Proteobacteria, Actinobacteria and Chloroflexi were related to N-cycling, and Actinobacteria and Nitrospirae were related to both N- and P-cycling. Proteobacteria and Chlorobi affected C-cycling and multiple functionality indexes in the abandoned land. We conclude that increasing land-use intensity changed the soil bacterial community structure and decreased bacterial interactions. However, increases in α-diversity at the rock-soil interface in cultivated soils indicated that major soil functions related to biogeochemical cycling were maintained within keystone taxa in this microenvironment. Our study provides foundations to test the success of different regeneration practices in restoring soil microbial diversity and the multifunctionality of karst ecosystems.

Leaf phosphorus content of Quercus wutaishanica increases with total soil potassium in the Loess Plateau.

Phosphorus (P) is arguably more limiting than nitrogen for forest ecosystems being free of disturbances for lengthy time periods. The elucidation of multivariate relationships between foliar P and its primary drivers for dominant species is an urgent issue and formidable challenge for ecologists. Our goal was to evaluate the effects of primary drivers on foliar P of Quercus wutaishanica, the dominant species in broadleaved deciduous forest at the Loess Plateau, China. We sampled the leaves of 90 Q. wutaishanica individuals across broad climate and soil nutrient gradients at the Loess Plateau, China, and employed structural equation models (SEM) to evaluate multiple causal pathways and the relative importance of the drivers for foliar P per unit mass (Pmass) and per unit area (Parea). Our SEMs explained 73% and 81% of the variations in Pmass and Parea, respectively. Pmass was negatively correlated to leaf mass per area, positively correlated to leaf area, and increased with mean annual precipitation and total soil potassium. Parea was positively correlated to leaf mass per area, leaf dry weight, and increased significantly with total soil potassium. Our results demonstrated that leaf P content of Q. wutaishanica increased with total soil potassium in the Loess Plateau accordingly.

Determinants of the N content of Quercus wutaishanica leaves in the Loess Plateau: a structural equation modeling approach.

Most terrestrial ecosystems are nitrogen (N)-limited. The elucidation of the multivariate relationships among environmental drivers, leaf morphological traits, and foliar N of dominant species which are critical to the functioning of forests remains a critical challenge for ecologists. We sampled leaves of Quercus wutaishanica across a broad natural gradient in the Loess Plateau, China, and employed structural equation modelling to evaluate the causal pathways and the relative importance of drivers of the foliar N per unit area (Narea) and per unit mass (Nmass). We found that (1) Nmass and Narea were primarily affected by leaf morphological traits instead of environmental variables and that leaf morphological traits accounted for most of their variations; (2) the total soil potassium and phosphorus and mean annual precipitation had different effects on Nmass and Narea via different pathways and path coefficients, whereas the mean annual temperature and total soil N had non-significant effects on Nmass and Narea. Our results demonstrated that variations in Nmass and Narea within Quercus wutaishanica were strongly linked to their leaf morphological traits and that the leaf N was also influenced by mean annual precipitation and soil phosphorus and potassium instead of soil N in the Loess Plateau, China.