Isoprene emissions from plants are mediated by atmospheric CO2 concentrations

The tropical African tree species Acacia nigrescens Oliv. was grown in environmentally controlled growth chambers at three CO₂ concentrations representative of the Last Glacial Maximum (～180 ppmv), the present day (～380 ppmv), and likely mid‐21st century (～600 ppmv) CO₂ concentrations. Isoprene (C₅H₈) emissions, per unit leaf area, were greater at lower‐than‐current CO₂ levels and lower at higher‐than‐current CO₂ levels relative to controls grown at 380 ppmv CO₂. Changes in substrate availability and isoprene synthase (IspS) activity were identified as the mechanisms behind the observed leaf‐level emission response. In contrast, canopy‐scale emissions remained unaltered between the treatments as changes in leaf‐level emissions were offset by changes in biomass and leaf area. Substrate concentration and IspS activity‐CO₂ responses were used in a biochemical model, coupled to existing isoprene emission algorithms, to model isoprene emissions from A. nigrescens grown for over 2 years at three different CO₂ concentrations. The addition of the biochemical model allowed for the use of emission factors measured under present day CO₂ concentrations across all three CO₂ treatments. When isoprene emissions were measured from A. nigrescens in response to instantaneous changes in CO₂ concentration, the biochemical model satisfactorily represented the observed response. Therefore, the effect of changes in atmospheric CO₂ concentration on isoprene emission at any timescale can be modelled and predicted.     

Isoprene emission by plants is affected by transmissible wound signals

Isoprene (2-methyl 1,3-butadiene) is emitted from many plants, but the signals regulating isoprene emission are unknown. Mounting leaves in a gas exchange chamber or taking small leaf punches for biochemical analysis was found to reduce the rate of isoprene emission (Loreto & Sharkey 1993). This phenomenon was investigated by putting terminal leaflets of velvet bean (Mucuna deeringeniana L.) and kudzu [Pueraria lobaia (Willd) Ohwi.] into a gas exchange chamber and monitoring isoprene emission and photosynthesis. Lateral leaflets or remote leaves were then wounded or mechanically stimulated. The rate of isoprene emission was reduced after 1 min by up to 75% by burning a lateral leaflet with a match. Even a 7 ms^?1 (25km h^?1) wind imposed on a lateral leaflet reduced isoprene emission from the terminal leaflet by 18%. Photosynthesis rates were either unaffected by these treatments or reduced more slowly than isoprene emission rates, indicating that the effect of isoprene emission rates was not a consequence of changes in photosynthetic activity. Isoprene emission from a terminal leaflet was reduced by burning leaves above and below the monitored leaflet when on the same stem. The effect was much reduced if the burned leaf (all three leaflets) was on a different stem from the monitored leaflet. Reduction of the rate of isoprene emission was observed even when the burned leaf was 52 cm distant from the measured leaflet. Increasing the distance between the stressed leaf and the monitored leaf caused the effect to be slower and smaller. It is speculated that a signal is generated by wounding which propagates through the plant at 1.3 mm s^?1. This velocity was consistent throughout the measurements and is similar to the rate of propagation of electrical signals such as action potentials and variation potentials. The effect of the environmental stress, and particularly the wind effect, can be frequent in nature and should be considered when estimating local and regional emission of isoprene for modelling atmospheric chemistry. If leaf samples used for isoprene determination are exposed to the type of stress we investigated, isoprene emission inventories based on leaf level measurements will be underestimated.     

Isoprene emission from a subarctic peatland under enhanced UV-B radiation

Isoprene is a reactive hydrocarbon with an important role in atmospheric chemistry, and emissions from vegetation contribute to atmospheric carbon fluxes. The magnitude of isoprene emissions from arctic peatlands is not known, and it may be altered by increasing UV-B radiation. Isoprene emission was measured with the dynamic chamber method from a subarctic peatland under long-term enhancement of UV-B radiation targeted to correspond to a 20% loss in the stratospheric ozone layer. The site type of the peatland was a flark fen dominated by the moss Warnstorfia exannulata and sedges Eriophorum russeolum and Carex limosa. The relationship between species densities and the emission was also assessed. Isoprene emissions were significantly increased by enhanced UV-B radiation during the second (2004) and the fourth (2006) growing seasons under the UV-B exposure. Emissions were related to the density of E. russeolum. The dominant moss, W. exannulata, proved to emit small amounts of isoprene in a laboratory trial. Subarctic fens, even without Sphagnum moss, are a significant source of isoprene to the atmosphere, especially under periods of warm weather. Warming of the Arctic together with enhanced UV-B radiation may substantially increase the emissions.     

Isoprene synthase activity and its relation to isoprene emission in Quercus robur L. leaves

Pedunculate oak ( Quercus robur L.) is known as a strong isoprene (2-methyl-1,3-butadiene) emitter. Diurnal changes in isoprene emission were determined by branch enclosure measurements. In contrast to the diurnal cycle in emission rates, specific isoprene synthase activity in the leaves remained unchanged. Based on in vitro enzyme activity and its temperature dependency, an isoprene synthesis capacity at specific leaf temperatures was calculated. The comparison of these 'leaf temperature-dependent enzyme capacities' and the measured emission rates revealed that the enzyme activity of isoprene synthase is comparable to the observed isoprene emission rates. In addition, variation in the isoprene synthase activity of the leaves due to changes in light intensity during leaf development was investigated. A 50% reduction of light intensity by shading of single branches reduced isoprene synthase activity by ≈ 60% compared with full sunlight. The calculation of isoprene synthesis capacities based on enzymatic data obtained under optimum reaction conditions, corrected for actual leaf temperature and related to leaf surface area, provides a sound basis for predicting the isoprene emission potential of plants.     

Isoprene emission and primary metabolism in Phragmites australis grown under different phosphorus levels

Aquatic plants are generally used for wastewater purification and phytoremediation, but some of them also emit large amounts of isoprene, the most abundant biogenic volatile organic compound. Since isoprenoid biosynthesis requires high amounts of phosphorylated intermediates, the emission may also be controlled by inorganic phosphorus concentration (Pi) in leaves. We carried out experiments to determine the emission of isoprene from Phragmites australis plants used in reconstructed wetlands to phytoremediate elevated levels of phosphorus contributed by urban wastes. Four groups of plants were grown hydroponically in water containing different levels of KH(2)PO(4). High levels of phosphorus in the water resulted in high Pi in the leaves. High Pi stimulated photosynthesis at intercellular CO(2) concentrations lower and higher than ambient, implying higher ribulose 1,5-bisphosphate carboxylase (Rubisco) activity and higher ribulose 1,5-bisphosphate regeneration rates, respectively. However, isoprene emission was substantially lower at high Pi than at low Pi, and was not associated to photosynthesis rates at high Pi. This surprising result suggests that isoprene is limited by processes other than photosynthetic intermediate availability or by energetic (ATP) requirements under high Pi levels. Irrespective of the mechanism responsible for the observed reduction of isoprene emission, our results show that Phragmites plants may effectively remove phosphorus from water without concurrently increase isoprene emission, at least on a leaf area basis. Thus, Phragmites used in reconstructed wetlands for phytoremediation of urban wastes rich of phosphates will not contribute high loads of hydrocarbons which may influence air quality over urban and peri-urban areas.     

The Influence of Light and Temperature on Isoprene Emission Rates from Live Oak

There is a growing awareness of the role of vegetation as a source of reactive hydrocarbons that may serve as photochemical oxidant precursors. A study was designed to assess independently the influence of variable light and temperature on isoprene emissions from live oak (Quercus virginiana Mill.). Plants were conditioned in a growth chamber and then transferred to an environmentally controlled gas-exchange chamber. Samples of the chamber atmosphere were collected; isoprene was concentrated cryogenically and measured by gas chromatography. A logistic function was used to model isoprene emission rates. Under regimes of low temperature (20°C) or darkness, isoprene emissions were lowest. With increasing temperature or light intensity, the rate of isoprene emission increased, reaching maxima at 800 μE m^-2 s^-1 and 40–44°C, respectively. Higher temperatures caused a large decrease in emissions. Since the emissions of isoprene were light-saturated at moderate intensities, temperature appeared to be the main factor controlling emissions during most of the day. Carbon lost through isoprene emissions accounted for 0.1 to 2% of the carbon fixed during photosynthesis depending on light intensity and temperature.     

Supply of precursors for carotenoid biosynthesis in plants

Carotenoids are isoprenoids of industrial and nutritional interest produced by all photosynthetic organisms, including plants. Too often, the metabolic engineering of plant carotenogenesis has been obstructed by our limited knowledge on how the endogenous pathway interacts with other related metabolic pathways, particularly with those involved in the production of isoprenoid precursors. However, recent discoveries are providing new insights into this field. All isoprenoids derive from prenyl diphosphate precursors. In the case of carotenoids, these precursors are produced predominantly by the methylerythritol 4-phosphate (MEP) pathway in plants. This review focuses on the progress in our understanding of how manipulation of the MEP pathway impacts carotenoid biosynthesis and on the discoveries underlining the central importance of coordinating the supply of MEP-derived precursors with the biosynthesis of carotenoids and other derived isoprenoids.     

Contribution of hydroxymethylbutenyl diphosphate synthase to carotenoid biosynthesis in bacteria and plants

The methylerythritol 4-phosphate (MEP) pathway synthesizes the precursors of carotenoids and other isoprenoids in bacteria and plant plastids. Despite recent progress in the identification of rate-determining steps, the relative contribution of most pathway enzymes to flux control remains to be established. In this work we investigated whether upregulated levels of hydroxymethylbutenyl diphosphate synthase (HDS) could increase the metabolic flux through this pathway, as judged by endpoint (carotenoid) measurements. Unlike other MEP pathway enzymes, however, increasing the levels of an active HDS protein in carotenoid-producing Escherichia coli cells and transgenic Arabidopsis thaliana plants did not result in an enhanced accumulation of MEP-derived isoprenoids. Our data suggest that enhanced flux through the MEP pathway for peak demand periods in bacteria and plastids does not require increased HDS activity.     

Longevity of clonal plants: why it matters and how to measure it

Background

Species'' life-history and population dynamics are strongly shaped by the longevity of individuals, but life span is one of the least accessible demographic traits, particularly in clonal plants. Continuous vegetative reproduction of genets enables persistence despite low or no sexual reproduction, affecting genet turnover rates and population stability. Therefore, the longevity of clonal plants is of considerable biological interest, but remains relatively poorly known.

Scope

Here, we critically review the present knowledge on the longevity of clonal plants and discuss its importance for population persistence. Direct life-span measurements such as growth-ring analysis in woody plants are relatively easy to take, although, for many clonal plants, these methods are not adequate due to the variable growth pattern of ramets and difficult genet identification. Recently, indirect methods have been introduced in which genet size and annual shoot increments are used to estimate genet age. These methods, often based on molecular techniques, allow the investigation of genet size and age structure of whole populations, a crucial issue for understanding their viability and persistence. However, indirect estimates of clonal longevity are impeded because the process of ageing in clonal plants is still poorly understood and because their size and age are not always well correlated. Alternative estimators for genet life span such as somatic mutations have recently been suggested.

Conclusions

Empirical knowledge on the longevity of clonal species has increased considerably in the last few years. Maximum age estimates are an indicator of population persistence, but are not sufficient to evaluate turnover rates and the ability of long-lived clonal plants to enhance community stability and ecosystem resilience. In order to understand the dynamics of populations it will be necessary to measure genet size and age structure, not only life spans of single individuals, and to use such data for modelling of genet dynamics.     

Isolation and Purification of Protein Responsible for the Conversion of Dimethylallylpyrophosphate from Poplar Leaves into Isoprene

The protein converting dimethylallylpyrophosphate (DMAPP) into isoprene in vitrowas isolated and purified 3000-fold from leaves of berry-bearing poplar (Populus deltoidesMarsh.). As the enzyme was purified, its specific activity increased and at the final stage reached 266 nmol/(min mg protein). The enzyme was eluted by anion-exchange chromatography in a 120–170 mM NaCl gradient and by chromatography on the hydroxyapatite column in 170 mM sodium phosphate. The active molecular weight of the protein determined by gel filtration was 100–110 kD. As the enzyme was purified, the K _Mvalue increased from 2 to 9 mM. A parallelism isoprene emission from DMAPP and an increase in the specific activity of the enzyme as it was purified proved that the enzyme catalyzed isoprene emission.     

大气二氧化碳浓度增加对木本植物BVOCs释放的影响

植物挥发性有机化合物(biogenic volatile organic compounds,BVOCs)在近地表臭氧和二次有机气溶胶生成中有重要作用,而大气CO2浓度上升对植物BVOCs释放有显著影响。利用Meta-analysis方法对已发表的数据进行整合分析发现:(1)总体而言,大气CO2浓度增加会导致不同木本植物(常绿与落叶) BVOCs释放降低;(2)就不同木本植物BVOCs释放而言,大气CO2浓度增加主要导致落叶植物BVOCs释放速率降低,而常绿植物则以增加为主;(3)就植物释放BVOCs种类而言,大气CO2浓度增加显著降低异戊二烯的释放速率,对单萜烯释放速率则无显著影响。结果可为阐明陆地生态系统BVOCs释放对全球CO2浓度增加的响应提供依据。     

1-Deoxy-D-xylulose 5-phosphate reductoisomerase: an overview

The methylerythritol phosphate pathway to isoprenoids, an alternate biosynthetic route present in many bacteria, algae, plants, and the malarial parasite Plasmodium falciparum, has become an attractive target for the development of new antimalarial and antibacterial compounds. The second enzyme in this pathway, 1-deoxy-D-xylulose 5-phosphate reductoisomerase (DXR; EC 1.1.1.267), has been shown to be the molecular target for fosmidomycin, a promising antimalarial drug. This enzyme converts 1-deoxy-D-xylulose 5-phosphate (DXP) into the branched compound 2-C-methyl-D-erythritol 4-phosphate (MEP). The transformation of DXP into MEP requires an isomerization, followed by a NADPH-dependent reduction. The discovery of DXR, its subsequent characterization, and the identification of inhibitors will be presented.     

Isoprene emission from aspen leaves : influence of environment and relation to photosynthesis and photorespiration

Isoprene emission rates from quaking aspen (Populus tremuloides Michx.) leaves were measured simultaneously with photosynthesis rate, stomatal conductance, and intercellular CO₂ partial pressure. Isoprene emission required the presence of CO₂ or O₂, but not both. The light response of isoprene emission rate paralleled that of photosynthesis. Isoprene emission was inhibited by decreasing ambient O₂ from 21% to 2%, only when there was oxygen insensitive photosynthesis. Mannose (10 millimolar) fed through cut stems resulted in strong inhibition of isoprene emission rate and is interpreted as evidence that isoprene biosynthesis requires either the export of triose phosphates from the chloroplast, or the continued synthesis of ATP. Light response experiments suggest that photosynthetically generated reductant or ATP is required for isoprene biosynthesis. Isoprene biosynthesis and emission are not directly linked to glycolate production through photorespiration, contrary to previous reports. Isoprene emission rate was inhibited by above-ambient CO₂ partial pressures (640 microbar outside and 425 microbar inside the leaf). The inhibition was not due to stomatal closure. This was established by varying ambient humidity at normal and elevated CO₂ partial pressures to measure isoprene emission rates over a range of stomatal conductances. Isoprene emission rates were inhibited at elevated CO₂ despite no change in stomatal conductance. Addition of abscisic acid to the transpiration stream dramatically inhibited stomatal conductance and photosynthesis rate, with a slight increase in isoprene emission rate. Thus, isoprene emission is independent of stomatal conductance, and may occur through the cuticle. Temperature had an influence on isoprene emission rate, with the Q₁₀ being 1.8 to 2.4 between 35 and 45°C. At these high temperatures the amount of carbon lost through isoprene emission was between 2.5 and 8% of that assimilated through photosynthesis. This represents a significant carbon cost that should be taken into account in determining midsummer carbon budgets for plants that are isoprene emitters.     

Ca transfer from the ER to mitochondria: When, how and why

The heterogenous subcellular distribution of a wide array of channels, pumps and exchangers allows extracellular stimuli to induce increases in cytoplasmic Ca²⁺ concentration ([Ca²⁺]_c) with highly defined spatial and temporal patterns, that in turn induce specific cellular responses (e.g. contraction, secretion, proliferation or cell death). In this extreme complexity, the role of mitochondria was considered marginal, till the direct measurement with targeted indicators allowed to appreciate that rapid and large increases of the [Ca²⁺] in the mitochondrial matrix ([Ca²⁺]_m) invariably follow the cytosolic rises. Given the low affinity of the mitochondrial Ca²⁺ transporters, the close proximity to the endoplasmic reticulum (ER) Ca²⁺-releasing channels was shown to be responsible for the prompt responsiveness of mitochondria. In this review, we will summarize the current knowledge of: i) the mitochondrial and ER Ca²⁺ channels mediating the ion transfer, ii) the structural and molecular foundations of the signaling contacts between the two organelles, iii) the functional consequences of the [Ca²⁺]_m increases, and iv) the effects of oncogene-mediated signals on mitochondrial Ca²⁺ homeostasis. Despite the rapid progress carried out in the latest years, a deeper molecular understanding is still needed to unlock the secrets of Ca²⁺ signaling machinery.     

A model of isoprene emission based on energetic requirements for isoprene synthesis and leaf photosynthetic properties for Liquidambar and Quercus

We present a physiological model of isoprene (2-methyl-1,3-butadiene) emission which considers the cost for isoprene synthesis, and the production of reductive equivalents in reactions of photosynthetic electron transport for Liquidambar styraciflua L. and for North American and European deciduous temperate Quercus species. In the model, we differentiate between leaf morphology (leaf dry mass per area, M_A, g m ^{? 2}) altering the content of enzymes of isoprene synthesis pathway per unit leaf area, and biochemical potentials of average leaf cells determining their capacity for isoprene emission. Isoprene emission rate per unit leaf area ( μ mol m ^{? 2} s ^{? 1}) is calculated as the product of M_A, the fraction of total electron flow used for isoprene synthesis ( ? , mol mol ^{? 1}), the rate of photosynthetic electron transport (J) per unit leaf dry mass (J_m, μ mol g ^{? 1} s ^{? 1}), and the reciprocal of the electron cost of isoprene synthesis [mol isoprene (mol electrons ^{? 1})]. The initial estimate of electron cost of isoprene synthesis is calculated according to the 1-deoxy- D -xylulose-5-phosphate pathway recently discovered in the chloroplasts, and is further modified to account for extra electron requirements because of photorespiration. The rate of photosynthetic electron transport is calculated by a process-based leaf photosynthesis model. A satisfactory fit to the light-dependence of isoprene emission is obtained using the light response curve of J, and a single value of ? , that is dependent on the isoprene synthase activity in the leaves. Temperature dependence of isoprene emission is obtained by combining the temperature response curves of photosynthetic electron transport, the shape of which is related to long-term temperature during leaf growth and development, and the specific activity of isoprene synthase, which is considered as essentially constant for all plants. The results of simulations demonstrate that the variety of temperature responses of isoprene emission observed within and among the species in previous studies may be explained by different optimum temperatures of J and/or limited maximum fraction of electrons used for isoprene synthesis. The model provides good fits to diurnal courses of field measurements of isoprene emission, and is also able to describe the changes in isoprene emission under stress conditions, for example, the decline in isoprene emission in water-stressed leaves.     

Cinderella story: PI4P goes from precursor to key signaling molecule

Abstract

Phosphatidylinositol lipids are signaling molecules involved in nearly all aspects of cellular regulation. Production of phosphatidylinositol 4-phosphate (PI4P) has long been recognized as one of the first steps in generating poly-phosphatidylinositol phosphates involved in actin organization, cell migration, and signal transduction. In addition, progress over the last decade has brought to light independent roles for PI4P in membrane trafficking and lipid homeostasis. Here, we describe recent advances that reveal the breadth of processes regulated by PI4P, the spectrum of PI4P effectors, and the mechanisms of spatiotemporal control that coordinate crosstalk between PI4P and cellular signaling pathways.     

Enhanced flux through the methylerythritol 4-phosphate pathway in Arabidopsis plants overexpressing deoxyxylulose 5-phosphate reductoisomerase

The methylerythritol 4-phosphate (MEP) pathway synthesizes the precursors for an astonishing diversity of plastid isoprenoids, including the major photosynthetic pigments chlorophylls and carotenoids. Since the identification of the first two enzymes of the pathway, deoxyxylulose 5-phoshate (DXP) synthase (DXS) and DXP reductoisomerase (DXR), they both were proposed as potential control points. Increased DXS activity has been shown to up-regulate the production of plastid isoprenoids in all systems tested, but the relative contribution of DXR to the supply of isoprenoid precursors is less clear. In this work, we have generated transgenic Arabidopsis thaliana plants with altered DXS and DXR enzyme levels, as estimated from their resistance to clomazone and fosmidomycin, respectively. The down-regulation of DXR resulted in variegation, reduced pigmentation and defects in chloroplast development, whereas DXR-overexpressing lines showed an increased accumulation of MEP- derived plastid isoprenoids such as chlorophylls, carotenoids, and taxadiene in transgenic plants engineered to produce this non-native isoprenoid. Changes in DXR levels in transgenic plants did not result in changes in␣DXS gene expression or enzyme accumulation, confirming that the observed effects on plastid isoprenoid levels in DXR-overexpressing lines were not an indirect consequence of altering DXS levels. The results indicate that the biosynthesis of MEP (the first committed intermediate of the pathway) limits the production of downstream isoprenoids in Arabidopsis chloroplasts, supporting a role for DXR in the control of the metabolic flux through the MEP pathway.