Interplay between edaphic and climatic factors unravels plant and microbial diversity along an altitudinal gradient.

Altitude influences biodiversity and physiochemical soil attributes in terrestrial ecosystems. It is of immense importance to know the patterns of how interactions among climatic and edaphic factors influence plant and microbial diversity in various ecosystems, particularly along the gradients. We hypothesize that altitudinal variation determines the distribution of plant and microbial species as well as their interactions. To test the hypothesis, different sites with variable altitudes were selected. Analyses of edaphic factors revealed significant (p < 0.001) effects of the altitude. Soil ammonium and nitrate were strongly affected by it contrary to potassium (K), soil organic matter and carbon. The response patterns of individual taxonomic groups differed across the altitudinal gradient. Plant species and soil fungal diversity increased with increasing altitude, while soil archaeal and bacterial diversity decreased with increasing altitude. Plant species richness showed significant positive and negative interactions with edaphic and climatic factors. Fungal species richness was also significantly influenced by the soil ammonium, nitrate, available phosphorus, available potassium, electrical conductivity, and the pH of the soil, but showed non-significant interactions with other edaphic factors. Similarly, soil variables had limited impact on soil bacterial and archaeal species richness along the altitude gradient. Proteobacteria, Ascomycota, and Thaumarchaeota dominate soil bacterial, fungal, and archaeal communities, with relative abundance of 27.4%, 70.56%, and 81.55%, respectively. Additionally, Cynodon dactylon is most abundant plant species, comprising 22.33% of the recorded plant taxa in various study sites. RDA revealed that these communities influenced by certain edaphic and climatic factors, e.g., Actinobacteria strongly respond to MAT, EC, and C/N ratio, Ascomycota and Basidiomycota show strong associations with EC and MAP, respectively. Thaumarcheota are linked to pH, and OM, while Cyperus rotundus are sensitive to AI and EC. In conclusion, the observed variations in microbial as well as plant species richness and changes in soil properties at different elevations provide valuable insights into the factors determining ecosystem stability and multifunctionality in different regions.

Variations and driving factors of leaf functional traits in the dominant desert plant species along an environmental gradient in the drylands of China.

Leaf functional traits (LFTs) of desert plants are responsive, adaptable and highly plastic to their environment. However, the macroscale variation in LFTs and driving factors underlying this variation remain unclear, especially for desert plants. Here, we measured eight LFTs, including leaf carbon concentration (LCC), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), specific leaf area (SLA), leaf dry matter content (LDMC), leaf mass per area (LMA), leaf thickness (LTH) and leaf tissue density (LTD) across 114 sites along environmental gradient in the drylands of China and in Guazhou Common Garden and evaluated the effect of environment and phylogeny on the LFTs. We noted that for all species, the mean values of LCC, LNC, LPC, SLA, LDMC, LMA, LTH and LTD were 384.62 mg g-1, 19.91 mg g-1, 1.12 mg g-1, 79.62 cm2 g-1, 0.74 g g-1, 237.39 g m-2, 0.38 mm and 0.91 g cm-3, respectively. LFTs exhibited significant geographical variations and the LNC, LMA and LTH in the plants of Guazhou Common Garden were significantly higher than the field sites in the drylands of China. LDMC and LTD of plants in Guazhou Common Garden were, however, considerably lower than those in the drylands of China. LCC, LPC, LTH and LTD differed significantly among different plant lifeforms, while LNC, SLA, LDMC and LMA didn't show significant variations. We found that the environmental variables explained higher spatial variations (3.6-66.3 %) in LFTs than the phylogeny (1.8-54.2 %). The LCC significantly increased, while LDMC and LTD decreased with increased temperature and reduced precipitation. LPC, LDMC, LMA, and LTD significantly increased, while SLA and LTH decreased with increased aridity. However, leaf elements were not significantly correlated with soil nutrients. The mean annual precipitation was a key factor controlling variations in LFTs at the macroscale in the drylands of China. These findings will provide new insights to better understand the response of LFTs and plants adaptation along environmental gradient in drylands, and will serve as a reference for studying biogeographic patterns of leaf traits.

Plant trait networks reveal adaptation strategies in the drylands of China.

BACKGROUND: Plants accomplish multiple functions by the interrelationships between functional traits. Clarifying the complex relationships between plant traits would enable us to better understand how plants employ different strategies to adapt to the environment. Although increasing attention is being paid to plant traits, few studies focused on the adaptation to aridity through the relationship among multiple traits. We established plant trait networks (PTNs) to explore the interdependence of sixteen plant traits across drylands. RESULTS: Our results revealed significant differences in PTNs among different plant life-forms and different levels of aridity. Trait relationships for woody plants were weaker, but were more modularized than for herbs. Woody plants were more connected in economic traits, whereas herbs were more connected in structural traits to reduce damage caused by drought. Furthermore, the correlations between traits were tighter with higher edge density in semi-arid than in arid regions, suggesting that resource sharing and trait coordination are more advantageous under low drought conditions. Importantly, our results demonstrated that stem phosphorus concentration (SPC) was a hub trait correlated with other traits across drylands. CONCLUSIONS: The results demonstrate that plants exhibited adaptations to the arid environment by adjusting trait modules through alternative strategies. PTNs provide a new insight into understanding the adaptation strategies of plants to drought stress based on the interdependence among plant functional traits.

Concentrations and bioconcentration factors of leaf microelements in response to environmental gradients in drylands of China.

Determining response patterns of plant leaf elements to environmental variables would be beneficial in understanding plant adaptive strategies and in predicting ecosystem biogeochemistry processes. Despite the vital role of microelements in life chemistry and ecosystem functioning, little is known about how plant microelement concentrations, especially their bioconcentration factors (BCFs, the ratio of plant to soil concentration of elements), respond to large-scale environmental gradients, such as aridity, soil properties and anthropogenic activities, in drylands. The aim of the present study was to fill this important gap. We determined leaf microelement BCFs by measuring the concentrations of Mn, Fe, Ni, Cu and Zn in soils from 33 sites and leaves of 111 plants from 67 species across the drylands of China. Leaf microelement concentrations were maintained within normal ranges to satisfy the basic requirements of plants, even in nutrient-poor soil. Aridity, soil organic carbon (SOC) and electrical conductivity (EC) had positive effects, while soil pH had a negative effect on leaf microelement concentrations. Except for Fe, aridity affected leaf microelement BCFs negatively and indirectly by increasing soil pH and SOC. Anthropogenic activities and soil clay contents had relatively weak impacts on both leaf microelement concentrations and BCFs. Moreover, leaf microelement concentrations and BCFs shifted with thresholds at 0.89 for aridity and 7.9 and 8.9 for soil pH. Woody plants were positive indicator species and herbaceous plants were mainly negative indicator species of leaf microelement concentrations and BCFs for aridity and soil pH. Our results suggest that increased aridity limits the absorption of microelements by plant leaves and enhances leaf microelement concentrations. The identification of indicator species for the response of plant microelements to aridity and key soil characteristics revealed that woody species in drylands were more tolerant to environmental changes than herbaceous species.

Plant-soil-microbe interactions in maintaining ecosystem stability and coordinated turnover under changing environmental conditions.

Ecosystem functions directly depend upon biophysical as well as biogeochemical reactions occurring at the soil-microbe-plant interface. Environment is considered as a major driver of any ecosystem and for the distributions of living organisms. Any changes in climate may potentially alter the composition of communities i.e., plants, soil microbes and the interactions between them. Since the impacts of global climate change are not short-term, it is indispensable to appraise its effects on different life forms including soil-microbe-plant interactions. This article highlights the crucial role that microbial communities play in interacting with plants under environmental disturbances, especially thermal and water stress. We reviewed that in response to the environmental changes, actions and reactions of plants and microbes vary markedly within an ecosystem. Changes in environment and climate like warming, CO2 elevation, and moisture deficiency impact plant and microbial performance, their diversity and ultimately community structure. Plant and soil feedbacks also affect interacting species and modify community composition. The interactive relationship between plants and soil microbes is critically important for structuring terrestrial ecosystems. The anticipated climate change is aggravating the living conditions for soil microbes and plants. The environmental insecurity and complications are not short-term and limited to any particular type of organism. We have appraised effects of climate change on the soil inhabiting microbes and plants in a broader prospect. This article highlights the unique qualities of tripartite interaction between plant-soil-microbe under climate change.

Effects of environmental factors on anthocyanin accumulation in the fruits of Lycium ruthenicum Murray across different desert grasslands.

Anthocyanins can help plants adapt and resist adverse environments and have important nutritional and medicinal effects on human beings. However, how environmental factors affect the anthocyanins accumulation of plants and how to improve the anthocyanins content of plants in different soils needs further exploration. Hence, this study aimed to investigate the effects of environmental factors on the accumulation of cyanidin, petunidin, malvidin, and delphinidin in the fruits of Lycium ruthenicum in sandy desert grassland (SS), gravel desert grassland (GD), and saline-alkali desert grassland (SD) in the lower reaches of the Shiyang River Basin. The variable importance screened the key environmental factors affecting anthocyanin accumulation in projection (VIP) and multiple stepwise regressions. The structural equation model (SEM) was established to understand how the climate and soil factors affect the total anthocyanin accumulation. For establishing soil nutrient optimization schemes by partial least squares regression (PLS) and the simplex algorithm used to improve the anthocyanin content in different types of desert grassland. In SS, electrical conductivity (EC) and microbial biomass carbon (SMBC) showed highly significant and positive effects on the content of total anthocyanin, cyanidin, and petunidin. In GD, soil moisture and microbial biomass nitrogen (SNBN) significantly negatively affected total anthocyanin content. In SD, catalase (CAT), phosphatase (PHO), and total potassium (TK) had the greatest impact on total anthocyanin content. It is indicated that the targeted improvement measures are necessary to increase anthocyanin content in the fruit of Lycium ruthenicum.

Continental-scale niche differentiation of dominant topsoil archaea in drylands.

Archaea represent a diverse group of microorganisms often associated with extreme environments. However, an integrated understanding of biogeographical patterns of the specialist Haloarchaea and the potential generalist ammonia-oxidizing archaea (AOA) across large-scale environmental gradients remains limited. We hypothesize that niche differentiation determines their distinct distributions along environmental gradients. To test the hypothesis, we use a continental-scale research network including 173 dryland sites across northern China. Our results demonstrate that Haloarchaea and AOA dominate topsoil archaeal communities. As hypothesized, Haloarchaea and AOA show strong niche differentiation associated with two ecosystem types mainly found in China's drylands (i.e. deserts vs. grasslands), and they differ in the degree of habitat specialization. The relative abundance and richness of Haloarchaea are higher in deserts due to specialization to relatively high soil salinity and extreme climates, while those of AOA are greater in grassland soils. Our results further indicate a divergence in ecological processes underlying the segregated distributions of Haloarchaea and AOA. Haloarchaea are governed primarily by environmental-based processes while the more generalist AOA are assembled mostly via spatial-based processes. Our findings add to existing knowledge of large-scale biogeography of topsoil archaea, advancing our predictive understanding on changes in topsoil archaeal communities in a drier world.

Phylogenetic independence in the variations in leaf functional traits among different plant life forms in an arid environment.

Leaf traits of global plants reveal the fundamental trade-offs in plant resource acquisition to conservation strategies. However, which leaf traits are consistent, converged, or diverged among herbs, shrubs, and subshrubs in an arid environment remains unclear. In the present study, we evaluated the trade-offs in six leaf functional traits (LFTs): leaf fresh mass (LFM), leaf dry mass (LDM), leaf dry matter content (LDMC), leaf area (LA), specific leaf area (SLA), and leaf thickness (LTh) of 37 desert plant species. LFTs differed between different plant life forms; LFM, LDM, and LA were slightly higher in herbs, LDMC and LTh in shrubs, and SLA in subshrubs. Conversely, the correlations among LFTs were inconsistent in different life forms, which may indicate their different adaptation strategies in an arid environment. Legumes and C3 plants exhibited slightly higher LDMC, LA, and SLA than non-legumes and C4 plants, whereas non-legumes and C4 plants showed higher (nonsignificant) LFM, LDM, and LTh than legumes and C3 plants. A significant phylogenetic signal (PS) and maximum K-value were found for SLA (K = 0.32). LFTs exhibited convergent and divergent variations among different life forms. However, these variations in LFTs were not influenced by phylogeny. Together, these findings increase our understanding of the variations in ecological adaptations of desert plants as well as adaption strategies of different life forms in an arid environment.

Effects of biotic and abiotic factors on forest biomass fractions.

The extent to which key factors at the global scale influence plant biomass allocation patterns remains unclear. Here, we provide a theory about how biotic and abiotic factors influence plant biomass allocation and evaluate its predictions using a large global database for forested communities. Our analyses confirm theoretical predictions that temperature, precipitation, and plant height and density jointly regulate the quotient of leaf biomass and total biomass, and that they have a much weaker effect on shoot (leaf plus stem) biomass fractions at a global scale. Moreover, biotic factors have larger effects than abiotic factors. Climatic variables act equally on shoot and root growth, and differences in plant body size and age, as well as community species composition, which vary with climate in ways that drown out the variations in biomass fractions. The theory and data presented here provide mechanistic explanations of why climate has little effect on biomass fractions.

Aridity-driven shift in biodiversity-soil multifunctionality relationships.

Relationships between biodiversity and multiple ecosystem functions (that is, ecosystem multifunctionality) are context-dependent. Both plant and soil microbial diversity have been reported to regulate ecosystem multifunctionality, but how their relative importance varies along environmental gradients remains poorly understood. Here, we relate plant and microbial diversity to soil multifunctionality across 130 dryland sites along a 4,000 km aridity gradient in northern China. Our results show a strong positive association between plant species richness and soil multifunctionality in less arid regions, whereas microbial diversity, in particular of fungi, is positively associated with multifunctionality in more arid regions. This shift in the relationships between plant or microbial diversity and soil multifunctionality occur at an aridity level of ∼0.8, the boundary between semiarid and arid climates, which is predicted to advance geographically ∼28% by the end of the current century. Our study highlights that biodiversity loss of plants and soil microorganisms may have especially strong consequences under low and high aridity conditions, respectively, which calls for climate-specific biodiversity conservation strategies to mitigate the effects of aridification.

Effects of Water and Energy on Plant Diversity along the Aridity Gradient across Dryland in China.

Plants need water and energy for their growth and reproduction. However, how water and energy availability influence dryland plant diversity along the aridity gradient in water-limited regions is still lacking. Hence, quantitative analyses were conducted to evaluate the relative importance of water and energy to dryland plant diversity based on 1039 quadrats across 184 sites in China's dryland. The results indicated that water availability and the water-energy interaction were pivotal to plant diversity in the entire dryland and consistent with the predictions of the water-energy dynamic hypothesis. The predominance of water limitation on dryland plant diversity showed a weak trend with decreasing aridity, while the effects of energy on plants were found to be significant in mesic regions. Moreover, the responses of different plant lifeforms to water and energy were found to vary along the aridity gradient. In conclusion, the study will enrich the limited knowledge about the effects of water and energy on plant diversity (overall plants and different lifeforms) in the dryland of China along the aridity gradient.

Convergent Variations in the Leaf Traits of Desert Plants.

Convergence is commonly caused by environmental filtering, severe climatic conditions and local disturbance. The basic aim of the present study was to understand the pattern of leaf traits across diverse desert plant species in a common garden, in addition to determining the effect of plant life forms (PLF), such as herb, shrub and subshrub, phylogeny and soil properties on leaf traits. Six leaf traits, namely carbon (C), nitrogen (N), phosphorus (P), potassium (K), δ13C and leaf water potential (LWP) of 37 dominant desert plant species were investigated and analyzed. The C, N, K and δ13C concentrations in leaves of shrubs were found higher than herbs and subshrubs; however, P and LWP levels were higher in the leaves of subshrubs following herbs and shrubs. Moreover, leaf C showed a significant positive correlation with N and a negative correlation with δ13C. Leaf N exhibited a positive correlation with P. The relationship between soil and plant macro-elements was found generally insignificant but soil C and N exhibited a significant positive correlation with leaf P. Taxonomy showed a stronger effect on leaf C, N, P and δ13C than soil properties, explaining >50% of the total variability. C3 plants showed higher leaf C, N, P, K and LWP concentration than C4 plants, whereas C4 plants had higher δ13C than C3 plants. Legumes exhibited higher leaf C, N, K and LWP than nonlegumes, while nonlegumes had higher P and δ13C concentration than legumes. In all the species, significant phylogenetic signals (PS) were detected for C and N and nonsignificant PS for the rest of the leaf traits. In addition, these phylogenetic signals were found lower (K-value < 1), and the maximum K-value was noted for C (K = 0.35). The plants of common garden evolved and adapted themselves for their survival in the arid environment and showed convergent variations in their leaf traits. However, these variations were not phylogenetics-specific. Furthermore, marks of convergence found in leaf traits of the study area were most likely due to the environmental factors.

Water content quantitatively affects metabolic rates over the course of plant ontogeny.

Plant metabolism determines the structure and dynamics of ecological systems across many different scales. The metabolic theory of ecology quantitatively predicts the scaling of metabolic rate as a function of body size and temperature. However, the role of tissue water content has been neglected even though hydration significantly affects metabolism, and thus ecosystem structure and functioning. Here, we use a general model based on biochemical kinetics to quantify the combined effects of water content, body size and temperature on plant metabolic rates. The model was tested using a comprehensive dataset from 205 species across 10 orders of magnitude in body size from seeds to mature large trees. We show that water content significantly influences mass-specific metabolic rates as predicted by the model. The scaling exponents of whole-plant metabolic rate vs body size numerically converge onto 1.0 after water content is corrected regardless of body size or ontogenetic stage. The model provides novel insights into how water content together with body size and temperature quantitatively influence plant growth and metabolism, community dynamics and ecosystem energetics.

A General Model for Seed and Seedling Respiratory Metabolism.

The ontogeny of seed plants usually involves a dormant dehydrated state and the breaking of dormancy and germination, which distinguishes it from that of most organisms. Seed germination and seedling establishment are critical ontogenetic stages in the plant life cycle, and both are fueled by respiratory metabolism. However, the scaling of metabolic rate with respect to individual traits remains poorly understood. Here, we tested metabolic scaling theory during seed germination and early establishment growth using a recently developed model and empirical data collected from 41 species. The results show that (i) the mass-specific respiration rate (Rm) was weakly correlated with body mass, mass-specific N content, and mass-specific C content; (ii) Rm conformed to a single Michaelis-Menten curve as a function of tissue water content; and (iii) the central parameters in the model were highly correlated with DNA content and critical enzyme activities. The model offers new insights and a more integrative scaling theory that quantifies the combined effects of tissue water content and body mass on respiratory metabolism during early plant ontogeny.

Global Data Analysis Shows That Soil Nutrient Levels Dominate Foliar Nutrient Resorption Efficiency in Herbaceous Species.

Nutrient resorption plays an important role in ecology because it has a profound effect on subsequent plant growth. However, our current knowledge about patterns of nutrient resorption, particularly among herbaceous species, at a global scale is still inadequate. Here, we present a meta-analysis using a global dataset of nitrogen (N) and phosphorus (P) resorption efficiency encompassing 227 perennial herbaceous species. This analysis shows that the N and P resorption efficiency (NRE and PRE, respectively), and N:P resorption ratios (NRE:PRE) across all herbaceous plant groups are 59.4, 67.5, and 0.89%, respectively. Across all species, NRE, PRE, and NRE:PRE, exhibited different patterns along climatic and soil nutrient gradients, i.e., NRE decreases with increasing mean annual precipitation (MAP) and soil N, PRE increases with aridity index (AI) but decreases with MAP and soil P, and NRE:PRE decreases with increasing potential evapotranspiration (PET), AI, and soil N:P. NRE, PRE, and NRE:PRE also differed in functional species group (graminoids vs. forbs). Soil nutrient level was the largest contributor to the total variations in NRE, PRE, and NRE:PRE, while climate and herbaceous types had relatively smaller effects on NRE, PRE, and NRE:PRE. Collectively, these trends can inform attempts to model biogeochemical cycling at a global scale.

A theoretical framework for whole-plant carbon assimilation efficiency based on metabolic scaling theory: a test case using Picea seedlings.

Simultaneous and accurate measurements of whole-plant instantaneous carbon-use efficiency (ICUE) and annual total carbon-use efficiency (TCUE) are difficult to make, especially for trees. One usually estimates ICUE based on the net photosynthetic rate or the assumed proportional relationship between growth efficiency and ICUE. However, thus far, protocols for easily estimating annual TCUE remain problematic. Here, we present a theoretical framework (based on the metabolic scaling theory) to predict whole-plant annual TCUE by directly measuring instantaneous net photosynthetic and respiratory rates. This framework makes four predictions, which were evaluated empirically using seedlings of nine Picea taxa: (i) the flux rates of CO(2) and energy will scale isometrically as a function of plant size, (ii) whole-plant net and gross photosynthetic rates and the net primary productivity will scale isometrically with respect to total leaf mass, (iii) these scaling relationships will be independent of ambient temperature and humidity fluctuations (as measured within an experimental chamber) regardless of the instantaneous net photosynthetic rate or dark respiratory rate, or overall growth rate and (iv) TCUE will scale isometrically with respect to instantaneous efficiency of carbon use (i.e., the latter can be used to predict the former) across diverse species. These predictions were experimentally verified. We also found that the ranking of the nine taxa based on net photosynthetic rates differed from ranking based on either ICUE or TCUE. In addition, the absolute values of ICUE and TCUE significantly differed among the nine taxa, with both ICUE and temperature-corrected ICUE being highest for Picea abies and lowest for Picea schrenkiana. Nevertheless, the data are consistent with the predictions of our general theoretical framework, which can be used to access annual carbon-use efficiency of different species at the level of an individual plant based on simple, direct measurements. Moreover, we believe that our approach provides a way to cope with the complexities of different ecosystems, provided that sufficient measurements are taken to calibrate our approach to that of the system being studied.

Models and tests of optimal density and maximal yield for crop plants.

We introduce a theoretical framework that predicts the optimum planting density and maximal yield for an annual crop plant. Two critical parameters determine the trajectory of plant growth and the optimal density, N(opt), where canopies of growing plants just come into contact, and competition: (i) maximal size at maturity, M(max), which differs among varieties due to artificial selection for different usable products; and (ii) intrinsic growth rate, g, which may vary with variety and environmental conditions. The model predicts (i) when planting density is less than N(opt), all plants of a crop mature at the same maximal size, M(max), and biomass yield per area increases linearly with density; and (ii) when planting density is greater than N(opt), size at maturity and yield decrease with -4/3 and -1/3 powers of density, respectively. Field data from China show that most annual crops, regardless of variety and life form, exhibit similar scaling relations, with maximal size at maturity, M(max), accounting for most of the variation in optimal density, maximal yield, and energy use per area. Crops provide elegantly simple empirical model systems to study basic processes that determine the performance of plants in agricultural and less managed ecosystems.

Insights into plant size-density relationships from models and agricultural crops.

There is general agreement that competition for resources results in a tradeoff between plant mass, M, and density, but the mathematical form of the resulting thinning relationship and the mechanisms that generate it are debated. Here, we evaluate two complementary models, one based on the space-filling properties of canopy geometry and the other on the metabolic basis of resource use. For densely packed stands, both models predict that density scales as M(-3/4), energy use as M(0), and total biomass as M(1/4). Compilation and analysis of data from 183 populations of herbaceous crop species, 473 stands of managed tree plantations, and 13 populations of bamboo gave four major results: (i) At low initial planting densities, crops grew at similar rates, did not come into contact, and attained similar mature sizes; (ii) at higher initial densities, crops grew until neighboring plants came into contact, growth ceased as a result of competition for limited resources, and a tradeoff between density and size resulted in critical density scaling as M(-0.78), total resource use as M(-0.02), and total biomass as M(0.22); (iii) these scaling exponents are very close to the predicted values of M(-3/4), M(0), and M(1/4), respectively, and significantly different from the exponents suggested by some earlier studies; and (iv) our data extend previously documented scaling relationships for trees in natural forests to small herbaceous annual crops. These results provide a quantitative, predictive framework with important implications for the basic and applied plant sciences.