Competitive performance of Pinus massoniana is related to scaling relationships at the individual plant and branch levels.

PREMISE: Competition is an important driver of tree mortality and thus affects forest structure and dynamics. Tree architectural traits, such as height-to-diameter (H-D) and branch length-to-diameter (L-d) relationships are thought to influence species competitiveness by affecting light capture. Unfortunately, little is known about how the H vs. D and L vs. d scaling exponents are related to tree performance (defined in the context of growth vigor) in competition. METHODS: Using data from field surveys of 1547 individuals and destructive sampling of 51 trees with 1086 first-order branches from a high-density Pinus massoniana forest, we explored whether the H vs. D and the L vs. d scaling exponents respectively differed numerically across tree performance and branch vertical position in crowns. RESULTS: The results indicated that (1) the H vs. D scaling exponent decreased as tree performance declined; (2) the L vs. d scaling exponent differed across tree performance classes (i.e., the scaling exponent of "inferior" trees was significantly larger than that of "moderate" and "superior" trees); (3) the L vs. d scaling exponent decreased as branch position approached ground level; and (4) overall, the branch scaling exponent decreased as tree performance improved in each crown layer, but decreased significantly in the intermediate layer. CONCLUSIONS: This study highlights the variation within (and linkage among) length-to-diameter scaling relationships across tree performance at the individual and branch levels. This linkage provides new insights into potential mechanisms of tree growth variation (and even further mortality) under competition in subtropical forests.

The complete mitochondrial genomes of Parabotia lijiangensis (Cypriniformes: Botiidae).

In this study, we obtained the 16,579 base pair (bp) mitochondrial DNA sequence of Parabotia lijiangensis. The mitogenome encodes 13 protein-coding genes, 22 tRNA genes, 2 rRNA genes, a control region, and has a nucleotide composition of A: 30.8%, T: 25.2%, G: 16.1%, and C: 27.9% (AT content: 56.0%). The complete mitogenome of P. lijiangensis provides essential and important DNA molecular data for further phylogenetic and evolutionary analysis of the Botiidae family.

Validation of an Enzyme-Driven Model Explaining Photosynthetic Rate Responses to Limited Nitrogen in Crop Plants.

The limited availability of nitrogen (N) is a fundamental challenge for many crop plants. We have hypothesized that the relative crop photosynthetic rate (P) is exponentially constrained by certain plant-specific enzyme activities, such as ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), NADP-glyceraldehyde-3-phosphate dehydrogenase (NADP-G3PDH), 3-phosphoglyceric acid (PGA) kinase, and chloroplast fructose-1,6-bisphosphatase (cpFBPase), in Triticum aestivum and Oryza sativa. We conducted a literature search to compile information from previous studies on C3 and C4 crop plants, to examine the photosynthetic rate responses to limited leaf [N] levels. We found that in Zea mays, NADP-malic enzyme (NADP-ME), PEP carboxykinase (PCK), and Rubisco activities were positively correlated with P. A positive correlation was also observed between both phosphoenolpyruvate carboxylase (PEPC) and Rubisco activity with leaf [N] in Sorghum bicolor. Key enzyme activities responded differently to P in C3 and C4 plants, suggesting that other factors, such as leaf [N] and the stage of leaf growth, also limited specific enzyme activities. The relationships followed the best fitting exponential relationships between key enzymes and the P rate in both C3 and C4 plants. It was found that C4 species absorbed less leaf [N] but had higher [N] assimilation rates (A rate) and higher maximum photosynthesis rates (Pmax ), i.e., they were able to utilize and invest more [N] to sustain higher carbon gains. All C3 species studied herein had higher [N] storage (Nstore) and higher absorption of [N], when compared with the C4 species. Nstore was the main [N] source used for maintaining photosynthetic capacity and leaf expansion. Of the nine C3 species assessed, rice had the greatest Pmax , thereby absorbing more leaf [N]. Elevated CO2 (eCO2) was also found to reduce the leaf [N] and Pmax in rice but enhanced the leaf [N] and N use efficiency of photosynthesis in maize. We concluded that eCO2 affects [N] allocation, which directly or indirectly affects Pmax . These results highlight the need to further study these physiological and biochemical processes, to better predict how crops will respond to eCO2 concentrations and limited [N].

An non-loglinear enzyme-driven law of photosynthetic scaling in two representative crop seedlings under different water conditions.

The loglinear pattern of respiratory scaling has been studied for over a century, while an increasing number of non-loglinear patterns have been found in the plant kingdom. Several previous studies had attempted to reconcile conflicting patterns from the aspects of statistical approaches and developmental stages of the organisms. However, the underlying enzymatic mechanism was largely ignored. Here, we propose an enzyme-driven law of photosynthetic scaling and test it in typical crop seedlings under different water conditions. The results showed that the key enzyme activity, the relative photosynthetic assimilation and the relative growth rate were all constrained by the available water, and the relationship between these biological traits and the available water supported our predictions. The enzyme-driven law appears to be more suitable to explain the curvature of photosynthetic scaling than the well-established power law, since it provides insight into the biochemical origin of photosynthetic assimilation.

Indirect effects of water availability in driving and predicting productivity in the Gobi desert.

Climate is the fundamental determinant of plant metabolism and net primary productivity (NPP). However, whether climate drives NPP directly or indirectly is not well understand. The Gobi desert across a precipitation gradient in the arid zone provides an ideal naturally-controlled platform for studying the precipitation-productivity relationships. We conducted 3-year experiments in four Gobi desert shrublands across an aridity gradient in Gansu Province of China to test the relationship between water availability and shrub productivity as well as the relative importance of the possible factors driving productivity (using piecewise structural equation modeling) and to explore the appropriate variables for predicting productivity (using three spatial models). The results showed that water availability indirectly affected the NPP via stand biomass, while stand biomass had a significant direct effect on NPP regardless of whether the leaf water content and stand height were considered. The model based on stand size (71.6%) and the model that contained both stand size and water availability (72.3%) explained more of the variation in the water-NPP relationships than the model based on water availability (37.3%). Our findings suggest that even in extremely water-limited areas, the effects of water availability on plant growth and the kinetics of plant metabolism could be indirect via plant size, demonstrating the importance of plant size as an indicator of shrub productivity. This study explains the mechanisms underlying the NPP driving pattern and proposes a practical NPP model for arid ecosystems.

Positive ecological effects of wind farms on vegetation in China's Gobi desert.

With the rapid development of wind power, there are increasing concerns about the negative ecological effects of its construction and operation. However, previous studies have mainly focused on the effects of wind farms on flying fauna (i.e., birds and bats) or climate change separately from communities or ecosystems, and little attention has been paid to vegetation during wind farm operation. Furthermore, few studies have referred to vulnerable ecosystems with low biomass and biodiversity. In this research, a field study was conducted to investigate the effects of wind farms on the individual traits, community structures and ecosystem functions of Gobi Desert ecosystems. The effects were measured by comparing interfering areas (IAs, located between 40 m and 90 m in the downstream direction of the wind turbine) with non-interfering areas (NIAs, located over 200 m from the wind turbine matrixes). The results showed that (1) plant individuals in IAs were less stressed and in better physiological states than those in NIAs; (2) for community structures, IA plants tended to be shorter and denser and had a higher coverage condition than that of NIA plants; and (3) ecosystem functions in IAs were significantly improved due to the existence of shrubs and higher biomass. Meanwhile, significant correlations were identified between the wind wake caused by the large spinning blades and the community structures. Constructing wind turbines in the Gobi Desert is a win-win strategy that both contributes to the growth of desert vegetation with a favourable microclimate and sufficiently utilizes wind power to produce clean energy.

Enzymatic origin and various curvatures of metabolic scaling in microbes.

The famous and controversial power law is a basal metabolic scaling model mainly derived from the "surface rule" or a fractal transport network. However, this law neglects biological mechanisms in the important active state. Here, we hypothesized that the relative metabolic rate and growth rate of actively growing microbes are driven by the changeable rate of their rate-limiting enzymes and concluded that natural logarithmic microbial metabolism (lnλ) and growth (or biomass) (lnM) are both dependent on limiting resources, and then developed novel models with interdependence between lnλ and lnM. We tested the models using the data obtained from the literature. We explain how and why the scaling is usually curved with the difference between microbial metabolic and growth (or biomass's) half-saturation constants (KM, Kλ) in the active state and agree that the linear relationship of the power law is a particular case under the given condition: KM = Kλ, which means that the enzyme dynamics may drive active and basal metabolic scaling relationships. Our interdependent model is more general than the power law, which is important for integrating the ecology and biochemical processes.

Glucose triggers stomatal closure mediated by basal signaling through HXK1 and PYR/RCAR receptors in Arabidopsis.

Sugars play important roles in regulating plant growth, development, and stomatal movement. Here, we found that glucose triggered stomatal closure in a dose- and time-dependent manner in Arabidopsis. Pharmacological data showed that glucose-induced stomatal closure was greatly inhibited by catalase [CAT; a reactive oxygen species (ROS) scavenger], diphenyleneiodonium chloride (DPI; an NADPH oxidase inhibitor), lanthanum chloride (LaCl3; a Ca2+ channel blocker), EGTA (a Ca2+ chelator), and two nitrate reductase (NR) inhibitors, tungstate and sodium azide (NaN3), while it was not affected by salicylhydroxamic acid (SHAM; a peroxidase inhibitor). Moreover, glucose induced ROS and nitric oxide (NO) production in guard cells of Arabidopsis. The ROS production was almost completely removed by CAT, strongly restricted by DPI, and was not affected by SHAM. NO production was partially suppressed by tungstate and NaN3, and the levels of NO were significantly reduced in the nia1-1nia2-5 mutant. Additionally, glucose-triggered stomatal closure was significantly impaired in gin1-1, gin2-1, pyr1pyl1pyl2pyl4, abi1-1, ost1, slac1-4, cpk6-1, and nia1-1nia2-5 mutants. Likewise, the reductions in leaf stomatal conductance (gs) and transpiration rate (E) caused by glucose were reversed in the above mutants. These results suggest that glucose-triggered stomatal closure may be dependent on basal signaling through PYR/RCAR receptors and hexokinase1 (HXK1).

Glucose- and mannose-induced stomatal closure is mediated by ROS production, Ca(2+) and water channel in Vicia faba.

Sugars act as vital signaling molecules that regulate plant growth, development and stress responses. However, the effects of sugars on stomatal movement have been unclear. In our study, we explored the effects of monosaccharides such as glucose and mannose on stomatal aperture. Here, we demonstrate that glucose and mannose trigger stomatal closure in a dose- and time-dependent manner in epidermal peels of broad bean (Vicia faba). Pharmacological studies revealed that glucose- and mannose-induced stomatal closure was almost completely inhibited by two reactive oxygen species (ROS) scavengers, catalase (CAT) and reduced glutathione (GSH), was significantly abolished by an NADPH oxidase inhibitor, diphenylene iodonium chloride (DPI), whereas they were hardly affected by a peroxidase inhibitor, salicylhydroxamic acid (SHAM). Furthermore, glucose- and mannose-induced stomatal closure was strongly inhibited by a Ca(2+) channel blocker, LaCl3 , a Ca(2+) chelator, ethyleneglycol-bis(beta-aminoethylether)-N,N'-tetraacetic acid (EGTA) and two water channel blockers, HgCl2 and dimethyl sulfoxide (DMSO); whereas the inhibitory effects of the water channel blockers were essentially abolished by the reversing agent ß-mercaptoethanol (ß-ME). These results suggest that ROS production mainly via NADPH oxidases, Ca(2+) and water channels are involved in glucose- and mannose-induced stomatal closure.

Plant Photosynthesis-Irradiance Curve Responses to Pollution Show Non-Competitive Inhibited Michaelis Kinetics.

Photosynthesis-irradiance (PI) curves are extensively used in field and laboratory research to evaluate the photon-use efficiency of plants. However, most existing models for PI curves focus on the relationship between the photosynthetic rate (Pn) and photosynthetically active radiation (PAR), and do not take account of the influence of environmental factors on the curve. In the present study, we used a new non-competitive inhibited Michaelis-Menten model (NIMM) to predict the co-variation of Pn, PAR, and the relative pollution index (I). We then evaluated the model with published data and our own experimental data. The results indicate that the Pn of plants decreased with increasing I in the environment and, as predicted, were all fitted well by the NIMM model. Therefore, our model provides a robust basis to evaluate and understand the influence of environmental pollution on plant photosynthesis.

Stomata prioritize their responses to multiple biotic and abiotic signal inputs.

Stomata are microscopic pores in leaf epidermis that regulate gas exchange between plants and the environment. Being natural openings on the leaf surface, stomata also serve as ports for the invasion of foliar pathogenic bacteria. Each stomatal pore is enclosed by a pair of guard cells that are able to sense a wide spectrum of biotic and abiotic stresses and respond by precisely adjusting the pore width. However, it is not clear whether stomatal responses to simultaneously imposed biotic and abiotic signals are mutually dependent on each other. Here we show that a genetically engineered Escherichia coli strain DH5α could trigger stomatal closure in Vicia faba, an innate immune response that might depend on NADPH oxidase-mediated ROS burst. DH5α-induced stomatal closure could be abolished or disguised under certain environmental conditions like low [CO2], darkness, and drought, etc. Foliar spraying of high concentrations of ABA could reduce stomatal aperture in high humidity-treated faba bean plants. Consistently, the aggressive multiplication of DH5α bacteria in Vicia faba leaves under high humidity could be alleviated by exogenous application of ABA. Our data suggest that a successful colonization of bacteria on the leaf surface is correlated with stomatal aperture regulation by a specific set of environmental factors.

Chlorella triggers stomatal closure mediated by NADPH oxidase and improves instantaneous water use efficiency in Vicia faba.

Stomatal closure induced by live microbes and microbe-associated molecular patterns (MAMPs) has been associated with early defense responses of plants. Chlorella is a unicellular autotrophic microorganism that can synthesize many bioactive substances with positive effects on humans, animals and plants. However, its effects on stomatal movement and instantaneous intrinsic water use efficiency (WUEi) in plants have been not explored yet. Our present work showed that application of Chlorella to isolated epidermal peels of Vicia faba induced stomatal closure in a dose-and time-dependent manner. Pharmacological study revealed that the Chlorella-triggered stomatal closure was mainly mediated by reactive oxygen species (ROS) production via NADPH oxidase. Additionally, exogenous application of optimal concentrations of Chlorella suspension caused an obvious reduction in leaf transpiration rate (E) without a parallel reduction in net photosynthetic rate (Pn), favoring the improvement of WUEi in Vicia faba. The chlorophyll fluorescence and content analysis further indicated that Chlorella had no effects on plant photosynthetic reactions center after short-term foliar application.

Chlorella induces stomatal closure via NADPH oxidase-dependent ROS production and its effects on instantaneous water use efficiency in Vicia faba.

Reactive oxygen species (ROS) have been established to participate in stomatal closure induced by live microbes and microbe-associated molecular patterns (MAMPs). Chlorella as a beneficial microorganism can be expected to trigger stomatal closure via ROS production. Here, we reported that Chlorella induced stomatal closure in a dose-and time-dependent manner in epidermal peels of Vicia faba. Using pharmacological methods in this work, we found that the Chlorella-induced stomatal closure was almost completely abolished by a hydrogen peroxide (H2O2) scavenger, catalase (CAT), significantly suppressed by an NADPH oxidase inhibitor, diphenylene iodonium chloride (DPI), and slightly affected by a peroxidase inhibitor, salicylhydroxamic acid (SHAM), suggesting that ROS production involved in Chlorella-induced stomatal closure is mainly mediated by DPI-sensitive NADPH oxidase. Additionally, Exogenous application of optimal concentrations of Chlorella suspension improved instantaneous water use efficiency (WUEi) in Vicia faba via a reduction in leaf transpiration rate (E) without a parallel reduction in net photosynthetic rate (Pn) assessed by gas-exchange measurements. The chlorophyll fluorescence and content analysis further demonstrated that short-term use of Chlorella did not influence plant photosynthetic reactions center. These results preliminarily reveal that Chlorella can trigger stomatal closure via NADPH oxidase-dependent ROS production in epidermal strips and improve WUEi in leave levels.

Scaling relationships between leaf mass and total plant mass across Chinese forests.

Biomass partitioning is important for illustrating terrestrial ecosystem carbon flux. West, Brown and Enquist (WBE) model predicts that an optimal 3/4 allometric scaling of leaf mass and total biomass of individual plants will be applied in diverse communities. However, amount of scientific evidence suggests an involvement of some biological and environmental factors in interpreting the variation of scaling exponent observed in empirical studies. In this paper, biomass information of 1175 forested communities in China was collected and categorized into groups in terms of leaf form and function, as well as their locations to test whether the allocation pattern was conserved or variable with internal and/or environmental variations. Model Type II regression protocol was adopted to perform all the regressions. The results empirically showed that the slopes varied significantly across diverse forested biomes, between conifer and broadleaved forests, and between evergreen and deciduous forests. Based on the results, leaf form and function and their relations to environments play a significant role in the modification of the WBE model to explore more accurate laws in nature.

The scaling relationships between leaf mass and leaf area of vascular plant species change with altitude.

The scaling relationship between leaf dry mass and leaf surface area has important implications for understanding the ability of plants to harvest sunlight and grow. Whether and how the scaling relationships vary across environmental gradients are poorly understood. We analyzed the scaling relationships between leaf mass and leaf area of 121 vascular plant species along an altitudinal gradient in a subtropical monsoon forest. The slopes increased significantly with altitude, it varied from less than 1 at low altitude to more than 1 at high altitude. This means that plants growing at high altitude allocate proportionately more biomass to support tissues in larger leaves and less in smaller leaves, whereas the reverse is true at low altitude. This pattern can be explained by different leaf strategies in response to environmental pressure and constrains.

Saccharomyces cerevisiae-induced stomatal closure mainly mediated by salicylhydroxamic acid-sensitive peroxidases in Vicia faba.

Saccharomyces cerevisiae induced stomatal closure in a dose-dependent manner on Vicia faba L. (cv. Daqingpi). Using pharmacological inhibitors in this study, we found that stomatal closure was completely inhibited by salicylhydroxamic acid (SHAM) and reduced glutathione (GSH), whereas slightly inhibited by diphenyleneiodonium chloride (DPI), suggesting that H2O2 was mostly produced by cell wall peroxidases. The specific NO scavenger (cPTIO), NO synthase (NOS) inhibitor NG-nitro-l-arginine methyl ester (l-NAME) and sodium azide (NaN3; inhibitor of nitrate reductase) prevented yeast-induced stomatal closure, suggesting that NO in guard cells of V. faba is derived from both NOS-like enzyme and nitrate reductase. Results of HgCl2 and ß-mercaptoethanol (ME) treatment (as a functional inhibitor of water channels and its reversing agent, respectively) suggest that water channels are involved in yeast-induced stomatal movements. CoCl2 (the blocker of calcium channel), LaCl3 (Ca(2+) antagonist) and EGTA (Ca(2+) chelator) also impaired yeast-induced stomatal closure. Thus, it is concluded that H2O2, NO, water channels and Ca(2+) are involved in yeast-induced stomatal closure.

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.

Sensitivity to abscisic acid modulates positive interactions between Arabidopsis thaliana individuals.

The ability of abscisic acid (ABA) to modulate positive interactions between Arabidopsis thaliana individuals under salinity stress was investigated using abi1-1 (insensitive to ABA), era1-2 (hypersensitive to ABA) mutant and wild type plants. The results showed that sensitivity to ABA affects relative interaction intensity (RII) between Arabidopsis thaliana individuals. The neighbor removal experiments also confirmed the role of phenotypic responses in linking plant-plant interactions and sensitivity to ABA. For abi1-1 mutants, the absolute value differences between neighbor removal and control of stem length, root length, leaf area, leaf thickness, flower density, above biomass/belowground biomass (A/U), photosynthetic rate, stomatal conductance, leaf water content and water-use efficiency were smaller than those of the wild type, while for era1-2 mutants, these absolute value differences were larger than those of the wild type. Thus, it is suggested that positive interactions between Arabidopsis thaliana individuals are at least partly modulated by different sensitivity to ABA through different physiological and phenotypic plasticity.

Scaling relationship between tree respiration rates and biomass.

The WBE theory proposed by West, Brown and Enquist predicts that larger plant respiration rate, R, scales to the three-quarters power of body size, M. However, studies on the R versus M relationship for larger plants (i.e. trees larger than saplings) have not been reported. Published respiration rates of field-grown trees (saplings and larger trees) were examined to test this relationship. Our results showed that for larger trees, aboveground respiration rates RA scaled as the 0.82-power of aboveground biomass MA, and that total respiration rates RT scaled as the 0.85-power of total biomass MT, both of which significantly deviated from the three-quarters scaling law predicted by the WBE theory, and which agreed with 0.81-0.84-power scaling of biomass to respiration across the full range of measured tree sizes for an independent dataset reported by Reich et al. (Reich et al. 2006 Nature 439, 457-461). By contrast, R scaled nearly isometrically with M in saplings. We contend that the scaling exponent of plant metabolism is close to unity for saplings and decreases (but is significantly larger than three-quarters) as trees grow, implying that there is no universal metabolic scaling in plants.