Variations of Leaf Cuticular Waxes Among C3 and C4 Gramineae Herbs.

PubMed

He, Yuji; Gao, Jianhua; Guo, Na; Guo, Yanjun

2016-11-01

Modern C4 plants are commonly distributed in hot and dry environments whereas C3 plants predominate in cool and shade areas. At the outmost of plant surface, the deposition and chemical composition of cuticular waxes vary under different environmental conditions. However, whether such variation of cuticular wax is related to the distribution of C3 and C4 under different environmental conditions is still not clear. In this study, leaves of six C3 Gramineae herbs distributed in spring, Roegneria kamoji, Polypogon fugax, Poa annua, Avena fatua, Alopecurus aequalis, and Oplismenus undulatifolius, and four C4 and one C3 Gramineae herbs distributed in summer, Digitaria sanguinalis, Eleusine indica, Setaria viridis, S. plicata, and O. undulatifolius, were sampled and analyzed for cuticular wax. Plates were the main epicuticular wax morphology in both C3 and C4 plants except S. plicata. The plates melted in C4 plants but not in C3 plants. The total cuticular wax amounts in C4 plants were significantly lower than those in C3 plants, except for O. undulatifolius. Primary alcohols were the most abundant compounds in C3 plants, whereas n-alkanes were relatively the most abundant compounds in C4 plants. C 29 was the most abundant n-alkane in C3 plants except for O. undulatifolius, whereas the most abundant n-alkane was C 31 or C 33 in C4 plants. The average chain length (ACL) of n-alkanes was higher in C4 than in C3 plants, whereas the ACL of n-alkanoic acids was higher in C3 than C4 plants. The cluster analysis based on the distribution of n-alkanes clearly distinguished C3 and C4 plants into two groups, except for O. undulatifolius which was grouped with C4 plants. These results suggest that the variations of cuticular waxes among C3 and C4 Gramineae herbs are related to the distribution of C3 and C4 plants under different environmental conditions. © 2016 Wiley-VHCA AG, Zurich, Switzerland.

Increasing water use efficiency along the C3 to C4 evolutionary pathway: a stomatal optimization perspective.

PubMed

Way, Danielle A; Katul, Gabriel G; Manzoni, Stefano; Vico, Giulia

2014-07-01

C4 photosynthesis evolved independently numerous times, probably in response to declining atmospheric CO2 concentrations, but also to high temperatures and aridity, which enhance water losses through transpiration. Here, the environmental factors controlling stomatal behaviour of leaf-level carbon and water exchange were examined across the evolutionary continuum from C3 to C4 photosynthesis at current (400 μmol mol(-1)) and low (280 μmol mol(-1)) atmospheric CO2 conditions. To this aim, a stomatal optimization model was further developed to describe the evolutionary continuum from C3 to C4 species within a unified framework. Data on C3, three categories of C3-C4 intermediates, and C4 Flaveria species were used to parameterize the stomatal model, including parameters for the marginal water use efficiency and the efficiency of the CO2-concentrating mechanism (or C4 pump); these two parameters are interpreted as traits reflecting the stomatal and photosynthetic adjustments during the C3 to C4 transformation. Neither the marginal water use efficiency nor the C4 pump strength changed significantly from C3 to early C3-C4 intermediate stages, but both traits significantly increased between early C3-C4 intermediates and the C4-like intermediates with an operational C4 cycle. At low CO2, net photosynthetic rates showed continuous increases from a C3 state, across the intermediates and towards C4 photosynthesis, but only C4-like intermediates and C4 species (with an operational C4 cycle) had higher water use efficiencies than C3 Flaveria. The results demonstrate that both the marginal water use efficiency and the C4 pump strength increase in C4 Flaveria to improve their photosynthesis and water use efficiency compared with C3 species. These findings emphasize that the advantage of the early intermediate stages is predominantly carbon based, not water related. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology.

Quo vadis C(4)? An ecophysiological perspective on global change and the future of C(4) plants.

PubMed

Sage, Rowan F; Kubien, David S

2003-01-01

C(4) plants are directly affected by all major global change parameters, often in a manner that is distinct from that of C(3) plants. Rising CO(2) generally stimulates C(3) photosynthesis more than C(4), but C(4) species still exhibit positive responses, particularly at elevated temperature and arid conditions where they are currently common. Acclimation of photosynthesis to high CO(2) occurs in both C(3) and C(4) plants, most notably in nutrient-limited situations. High CO(2) aggravates nitrogen limitations and in doing so may favor C(4) species, which have greater photosynthetic nitrogen use efficiency. C(4) photosynthesis is favored by high temperature, but global warming will not necessarily favor C(4) over C(3) plants because the timing of warming could be more critical than the warming itself. C(3) species will likely be favored where harsh winter climates are moderated, particularly where hot summers also become drier and less favorable to C(4) plant growth. Eutrophication of soils by nitrogen deposition generally favors C(3) species by offsetting the superior nitrogen use efficiency of C(4) species; this should allow C(3) species to expand at the expense of C(4) plants. Land-use change and biotic invasions are also important global change factors that affect the future of C(4) plants. Human exploitation of forested landscapes favors C(4) species at low latitude by removing woody competitors and opening gaps in which C(4) grasses can establish. Invasive C(4) grasses are causing widespread forest loss in Asia, the Americas and Oceania by accelerating fire cycles and reducing soil nutrient status. Once established, weedy C(4) grasses can prevent woodland establishment, and thus arrest ecological succession. In sum, in the future, certain C(4) plants will prosper at the expense of C(3) species, and should be able to adjust to the changes the future brings.

Contribution of the Alternative Respiratory Pathway to PSII Photoprotection in C3 and C4 Plants.

PubMed

Zhang, Zi-Shan; Liu, Mei-Jun; Scheibe, Renate; Selinski, Jennifer; Zhang, Li-Tao; Yang, Cheng; Meng, Xiang-Long; Gao, Hui-Yuan

2017-01-09

The mechanism by which the mitochondrial alternative oxidase (AOX) pathway contributes to photosystem II (PSII) photoprotection is in dispute. It was generally thought that the AOX pathway protects photosystems by dissipating excess reducing equivalents exported from chloroplasts through the malate/oxaloacetate (Mal/OAA) shuttle and thus preventing the over-reduction of chloroplasts. In this study, using the aox1a Arabidopsis mutant and nine other C3 and C4 plant species, we revealed an additional action model of the AOX pathway in PSII photoprotection. Although the AOX pathway contributes to PSII photoprotection in C3 leaves treated with high light, this contribution was observed to disappear when photorespiration was suppressed. Disruption or inhibition of the AOX pathway significantly decreased the photorespiration in C3 leaves. Moreover, the AOX pathway did not respond to high light and contributed little to PSII photoprotection in C4 leaves possessing a highly active Mal/OAA shuttle but with little photorespiration. These results demonstrate that the AOX pathway contributes to PSII photoprotection in C3 plants by maintaining photorespiration to detoxify glycolate and via the indirect export of excess reducing equivalents from chloroplasts by the Mal/OAA shuttle. This new action model explains why the AOX pathway does not contribute to PSII photoprotection in C4 plants. Copyright © 2017 The Author. Published by Elsevier Inc. All rights reserved.

Photosynthetic carbon metabolism in seagrasses C-labeling evidence for the c(3) pathway.

PubMed

Andrews, T J; Abel, K M

1979-04-01

The delta(13)C values of several seagrasses were considerably less negative than those of terrestrial C(3) plants and tended toward those of terrestrial C(4) plants. However, for Thalassia hemprichii (Ehrenb.) Aschers and Halophila spinulosa (R. Br.) Aschers, phosphoglycerate and other C(3) cycle intermediates predominated among the early labeled products of photosynthesis in (14)C-labeled seawater (more than 90% at the earliest times) and the labeling pattern at longer times was brought about by the operation of the C(3) pathway. Malate and aspartate together accounted for only a minor fraction of the total fixed label at all times and the kinetic data of this labeling were not at all consistent with these compounds being early intermediates in seagrass photosynthesis. Pulse-chase (14)C-labeling studies further substantiated these conclusions. Significant labeling of photorespiratory intermediates was observed in all experiments. The kinetics of total fixation of label during some steady-state and pulse-chase experiments suggested that there may be an intermediate pool of inorganic carbon of variable size closely associated with the leaves, either externally or internally. Such a pool may be one cause for the C(4)-like carbon isotope ratios of seagrasses.

Photosynthetic characteristics of an amphibious plant, Eleocharis vivipara: Expression of C4 and C3 modes in contrasting environments

PubMed Central

Ueno, Osamu; Samejima, Muneaki; Muto, Shoshi; Miyachi, Shigetoh

1988-01-01

Eleocharis vivipara Link, a freshwater amphibious leafless plant belonging to the Cyperaceae can grow in both terrestrial and submersed aquatic conditions. Two forms of E. vivipara obtained from these contrasting environments were examined for the characteristics associated with C4 and C3 photosynthesis. In the terrestrial form (δ 13C values = -13.5 to -15.4‰, where ‰ is parts per thousand), the culms, which are photosynthetic organs, possess a Kranz-type anatomy typical of C4 plants, and well-developed bundle-sheath cells contain numerous large chloroplasts. In the submersed form (δ 13C value = -25.9‰), the culms possess anatomical features characteristic of submersed aquatic plants, and the reduced bundle-sheath cells contain only a few small chloroplasts. 14C pulse-12C chase experiments showed that the terrestrial form and the submersed form fix carbon by way of the C4 pathway, with aspartate (40%) and malate (35%) as the main primary products, and by way of the C3 pathway, with 3-phosphoglyceric acid (53%) and sugar phosphates (14%) as the main primary products, respectively. The terrestrial form showed photosynthetic enzyme activities typical of the NAD-malic enzyme-C4 subtype, whereas the submersed form showed decreased activities of key C4 enzymes and an increased ribulose 1,5-bisphosphate carboxylase (EC 4.1.1.39) activity. These data suggest that this species can differentiate into the C4 mode under terrestrial conditions and into the C3 mode under submersed conditions. Images PMID:16593980

Climate-driven C4 plant distributions in China: divergence in C4 taxa

PubMed Central

Wang, Renzhong; Ma, Linna

2016-01-01

There have been debates on the driving factors of C4 plant expansion, such as PCO2 decline in the late Micocene and warmer climate and precipitation at large-scale modern ecosystems. These disputes are mainly due to the lack of direct evidence and extensive data analysis. Here we use mass flora data to explore the driving factors of C4 distribution and divergent patterns for different C4 taxa at continental scale in China. The results display that it is mean annual climate variables driving C4 distribution at present-day vegetation. Mean annual temperature is the critical restriction of total C4 plants and the precipitation gradients seem to have much less impact. Grass and sedge C4 plants are largely restricted to mean annual temperature and precipitation respectively, while Chenopod C4 plants are strongly restricted by aridity in China. Separate regression analysis can succeed to detect divergences of climate distribution patterns of C4 taxa at global scale. PMID:27302686

CO2 availability influences hydraulic function of C3 and C4 grass leaves

PubMed Central

Blackman, Chris J

2018-01-01

Abstract Atmospheric CO2 (ca) has increased since the last glacial period, increasing photosynthetic water use efficiency and improving plant productivity. Evolution of C4 photosynthesis at low ca led to decreased stomatal conductance (gs), which provided an advantage over C3 plants that may be reduced by rising ca. Using controlled environments, we determined how increasing ca affects C4 water use relative to C3 plants. Leaf gas exchange and mass per area (LMA) were measured for four C3 and four C4 annual, crop-related grasses at glacial (200 µmol mol−1), ambient (400 µmol mol−1), and super-ambient (640 µmol mol−1) ca. C4 plants had lower gs, which resulted in a water use efficiency advantage at all ca and was broadly consistent with slower stomatal responses to shade, indicating less pressure on leaf water status. At glacial ca, net CO2 assimilation and LMA were lower for C3 than for C4 leaves, and C3 and C4 grasses decreased leaf hydraulic conductance (Kleaf) similarly, but only C4 leaves decreased osmotic potential at turgor loss. Greater carbon availability in C4 leaves at glacial ca generated a different hydraulic adjustment relative to C3 plants. At current and future ca, C4 grasses have advantages over C3 grasses due to lower gs, lower stomatal sensitivity, and higher absolute water use efficiency. PMID:29538702

Compound-specific stable carbon isotopic signature of carbohydrate pyrolysis products from C3 and C4 plants.

PubMed

González-Pérez, José A; Jiménez-Morillo, Nicasio T; de la Rosa, José M; Almendros, Gonzalo; González-Vila, Francisco J

2016-02-01

Pyrolysis-compound specific isotopic analysis (Py-CSIA: Py-GC-(FID)-C-IRMS) is a relatively novel technique that allows on-line quantification of stable isotope proportions in chromatographically separated products released by pyrolysis. Validation of the Py-CSIA technique is compulsory for molecular traceability in basic and applied research. In this work, commercial sucrose from C4 (sugarcane) and C3 (sugarbeet) photosystem plants and admixtures were studied using analytical pyrolysis (Py-GC/MS), bulk δ(13)C IRMS and δ(13)C Py-CSIA. Major pyrolysis compounds were furfural (F), furfural-5-hydroxymethyl (HMF) and levoglucosan (LV). Bulk and main pyrolysis compound δ(13)C (‰) values were dependent on plant origin: C3 (F, -24.65 ± 0.89; HMF, -22.07 ± 0.41‰; LV, -21.74 ± 0.17‰) and C4 (F, -14.35 ± 0.89‰; HMF, -11.22 ± 0.54‰; LV, -11.44 ± 1.26‰). Significant regressions were obtained for δ(13)C of bulk and pyrolysis compounds in C3 and C4 admixtures. Furfural (F) was found (13)C depleted with respect to bulk and HMF and LV, indicating the incorporation of the light carbon atom in position 6 of carbohydrates in the furan ring after pyrolysis. This is the first detailed report on the δ(13)C signature of major pyrolytically generated carbohydrate-derived molecules. The information provided by Py-CSIA is valuable for identifying source marker compounds of use in food science/fraud detection or in environmental research. © 2015 Society of Chemical Industry.

Enhancement of growth, photosynthetic performance and yield by exclusion of ambient UV components in C3 and C4 plants.

PubMed

Kataria, Sunita; Guruprasad, K N; Ahuja, Sumedha; Singh, Bupinder

2013-10-05

A field experiment was conducted under tropical climate for assessing the effect of ambient UV-B and UV-A by exclusion of UV components on the growth, photosynthetic performance and yield of C3 (cotton, wheat) and C4 (amaranthus, sorghum) plants. The plants were grown in specially designed UV exclusion chambers, wrapped with filters that excluded UV-B (<315nm), UV-A+B (<400nm), transmitted all the UV (280-400nm) or without filters. All the four plant species responded to UV exclusion by a significant increase in plant height, leaf area, leaf biomass, total biomass accumulation and yield. Measurements of the chlorophyll, chlorophyll fluorescence parameters, gas exchange parameters and the activity of Ribulose-1,5-bisphosphate carboxylase (Rubisco) by fixation of (14)CO2 indicated a direct relationship between enhanced rate of photosynthesis and yield of the plants. Quantum yield of electron transport was enhanced by the exclusion of UV indicating better utilization of PAR assimilation and enhancement in reducing power in all the four plant species. Exclusion of UV-B in particular significantly enhanced the net photosynthetic rate, stomatal conductance and activity of Rubisco. Additional fixation of carbon due to exclusion of ambient UV-B was channeled towards yield as there was a decrease in the level of UV-B absorbing substances and an increase in soluble proteins in all the four plant species. The magnitude of the promotion in all the parameters studied was higher in dicots (cotton, amaranthus) compared to monocots (wheat, sorghum) after UV exclusion. The results indicated a suppressive action of ambient UV-B on growth and photosynthesis; dicots were more sensitive than monocots in this suppression while no great difference in sensitivity was found between C3 and C4 plants. Experiments indicated the suppressive action of ambient UV on carbon fixation and yield of C3 and C4 plants. Exclusion of solar UV-B will have agricultural benefits in both C3 and C4 plants

Engineering C4 photosynthesis into C3 chassis in the synthetic biology age.

PubMed

Schuler, Mara L; Mantegazza, Otho; Weber, Andreas P M

2016-07-01

C4 photosynthetic plants outperform C3 plants in hot and arid climates. By concentrating carbon dioxide around Rubisco C4 plants drastically reduce photorespiration. The frequency with which plants evolved C4 photosynthesis independently challenges researchers to unravel the genetic mechanisms underlying this convergent evolutionary switch. The conversion of C3 crops, such as rice, towards C4 photosynthesis is a long-standing goal. Nevertheless, at the present time, in the age of synthetic biology, this still remains a monumental task, partially because the C4 carbon-concentrating biochemical cycle spans two cell types and thus requires specialized anatomy. Here we review the advances in understanding the molecular basis and the evolution of the C4 trait, advances in the last decades that were driven by systems biology methods. In this review we emphasise essential genetic engineering tools needed to translate our theoretical knowledge into engineering approaches. With our current molecular understanding of the biochemical C4 pathway, we propose a simplified rational engineering model exclusively built with known C4 metabolic components. Moreover, we discuss an alternative approach to the progressing international engineering attempts that would combine targeted mutagenesis and directed evolution. © 2016 The Authors The Plant Journal © 2016 John Wiley & Sons Ltd.

From proto-Kranz to C4 Kranz: building the bridge to C4 photosynthesis.

PubMed

Sage, Rowan F; Khoshravesh, Roxana; Sage, Tammy L

2014-07-01

In this review, we examine how the specialized "Kranz" anatomy of C4 photosynthesis evolved from C3 ancestors. Kranz anatomy refers to the wreath-like structural traits that compartmentalize the biochemistry of C4 photosynthesis and enables the concentration of CO2 around Rubisco. A simplified version of Kranz anatomy is also present in the species that utilize C2 photosynthesis, where a photorespiratory glycine shuttle concentrates CO2 into an inner bundle-sheath-like compartment surrounding the vascular tissue. C2 Kranz is considered to be an intermediate stage in the evolutionary development of C4 Kranz, based on the intermediate branching position of C2 species in 14 evolutionary lineages of C4 photosynthesis. In the best-supported model of C4 evolution, Kranz anatomy in C2 species evolved from C3 ancestors with enlarged bundle sheath cells and high vein density. Four independent lineages have been identified where C3 sister species of C2 plants exhibit an increase in organelle numbers in the bundle sheath and enlarged bundle sheath cells. Notably, in all of these species, there is a pronounced shift of mitochondria to the inner bundle sheath wall, forming an incipient version of the C2 type of Kranz anatomy. This incipient version of C2 Kranz anatomy is termed proto-Kranz, and is proposed to scavenge photorespiratory CO2. By doing so, it may provide fitness benefits in hot environments, and thus represent a critical first stage of the evolution of both the C2 and C4 forms of Kranz anatomy. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.

C4GEM, a Genome-Scale Metabolic Model to Study C4 Plant Metabolism1[W][OA

PubMed Central

de Oliveira Dal’Molin, Cristiana Gomes; Quek, Lake-Ee; Palfreyman, Robin William; Brumbley, Stevens Michael; Nielsen, Lars Keld

2010-01-01

Leaves of C4 grasses (such as maize [Zea mays], sugarcane [Saccharum officinarum], and sorghum [Sorghum bicolor]) form a classical Kranz leaf anatomy. Unlike C3 plants, where photosynthetic CO2 fixation proceeds in the mesophyll (M), the fixation process in C4 plants is distributed between two cell types, the M cell and the bundle sheath (BS) cell. Here, we develop a C4 genome-scale model (C4GEM) for the investigation of flux distribution in M and BS cells during C4 photosynthesis. C4GEM, to our knowledge, is the first large-scale metabolic model that encapsulates metabolic interactions between two different cell types. C4GEM is based on the Arabidopsis (Arabidopsis thaliana) model (AraGEM) but has been extended by adding reactions and transporters responsible to represent three different C4 subtypes (NADP-ME [for malic enzyme], NAD-ME, and phosphoenolpyruvate carboxykinase). C4GEM has been validated for its ability to synthesize 47 biomass components and consists of 1,588 unique reactions, 1,755 metabolites, 83 interorganelle transporters, and 29 external transporters (including transport through plasmodesmata). Reactions in the common C4 model have been associated with well-annotated C4 species (NADP-ME subtypes): 3,557 genes in sorghum, 11,623 genes in maize, and 3,881 genes in sugarcane. The number of essential reactions not assigned to genes is 131, 135, and 156 in sorghum, maize, and sugarcane, respectively. Flux balance analysis was used to assess the metabolic activity in M and BS cells during C4 photosynthesis. Our simulations were consistent with chloroplast proteomic studies, and C4GEM predicted the classical C4 photosynthesis pathway and its major effect in organelle function in M and BS. The model also highlights differences in metabolic activities around photosystem I and photosystem II for three different C4 subtypes. Effects of CO2 leakage were also explored. C4GEM is a viable framework for in silico analysis of cell cooperation between M and BS

The C(4) plant lineages of planet Earth.

PubMed

Sage, Rowan F; Christin, Pascal-Antoine; Edwards, Erika J

2011-05-01

Using isotopic screens, phylogenetic assessments, and 45 years of physiological data, it is now possible to identify most of the evolutionary lineages expressing the C(4) photosynthetic pathway. Here, 62 recognizable lineages of C(4) photosynthesis are listed. Thirty-six lineages (60%) occur in the eudicots. Monocots account for 26 lineages, with a minimum of 18 lineages being present in the grass family and six in the sedge family. Species exhibiting the C(3)-C(4) intermediate type of photosynthesis correspond to 21 lineages. Of these, 9 are not immediately associated with any C(4) lineage, indicating that they did not share common C(3)-C(4) ancestors with C(4) species and are instead an independent line. The geographic centre of origin for 47 of the lineages could be estimated. These centres tend to cluster in areas corresponding to what are now arid to semi-arid regions of southwestern North America, south-central South America, central Asia, northeastern and southern Africa, and inland Australia. With 62 independent lineages, C(4) photosynthesis has to be considered one of the most convergent of the complex evolutionary phenomena on planet Earth, and is thus an outstanding system to study the mechanisms of evolutionary adaptation.

Photosynthetic diversity meets biodiversity: the C4 plant example.

PubMed

Sage, Rowan F; Stata, Matt

2015-01-01

Physiological diversification reflects adaptation for specific environmental challenges. As the major physiological process that provides plants with carbon and energy, photosynthesis is under strong evolutionary selection that gives rise to variability in nearly all parts of the photosynthetic apparatus. Here, we discuss how plants, notably those using C4 photosynthesis, diversified in response to environmental challenges imposed by declining atmospheric CO2 content in recent geological time. This reduction in atmospheric CO2 increases the rate of photorespiration and reduces photosynthetic efficiency. While plants have evolved numerous mechanisms to compensate for low CO2, the most effective are the carbon concentration mechanisms of C4, C2, and CAM photosynthesis; and the pumping of dissolved inorganic carbon, mainly by algae. C4 photosynthesis enables plants to dominate warm, dry and often salinized habitats, and to colonize areas that are too stressful for most plant groups. Because C4 lineages generally lack arborescence, they cannot form forests. Hence, where they predominate, C4 plants create a different landscape than would occur if C3 plants were to predominate. These landscapes (mostly grasslands and savannahs) present unique selection environments that promoted the diversification of animal guilds able to graze upon the C4 vegetation. Thus, the rise of C4 photosynthesis has made a significant contribution to the origin of numerous biomes in the modern biosphere. Copyright © 2014. Published by Elsevier GmbH.

In silico analysis of decomposed reflectances of C3 and C4 plants aiming at the effective assessment of crop needs

NASA Astrophysics Data System (ADS)

Baranoski, Gladimir V. G.; Van Leeuwen, Spencer; Chen, Tenn F.

2017-04-01

By separating the surface and subsurface components of foliar hyperspectral signatures using polarization optics, it is possible to enhance the remote discrimination of different plant species and optimize the assessment of different factors associated with their health status. These initiatives, in turn, can lead to higher crop yield and lower environmental impact. It is important to consider, however, that the main varieties of crops, represented by C3 (e.g., soy) and C4 (e.g., maize) plants, have markedly distinct morphological characteristics. Accordingly, the influence of these characteristics on their interactions with impinging light may affect the selection of optimal probe wavelengths for specific applications making use of combined hyperspectral and polarization measurements. In this paper, we compare the sensitivity of the total (including surface and subsurface components) and subsurface reflectance responses of C3 and C4 plants to different spectral and geometrical light incidence conditions. This investigation is supported by measured biophysical data and predictive light transport simulations. The results of our comparisons indicate that the total and subsurface reflectance responses of C3 and C4 plants depict well-defined patterns of sensitivity for varying illumination conditions. We believe that these patterns should be considered in the design of high-fidelity crop discrimination and monitoring procedures.

Effects of gaseous ammonia on intracellular pH values in leaves of C 3- and C 4-plants

NASA Astrophysics Data System (ADS)

Yin, Zu-Hua; Kaiser, Werner; Heber, Ulrich; Raven, John A.

Responses of cytosolic and vacuolar pH to different concentrations (1.3-5.4 μmol NH 3 mol -1 gas or 0.940-3.825 mg NH 3 m -3 gas) of gaseous NH 3 were studied in experiments of 3 h duration by recording changes in fluorescence of pyranine and esculin in leaves of C 3 and C 4 plants. After a lag phase of 0.5-4 min, the uptake of NH 3 at 50-200 nmol m -2 leaf area s -1 increased pyranine fluorescence, indicating cytosolic alkalinization in leaves of Pelargonium zonale L. (C 3) and Amaranthus caudatus L. (C 4). A smaller increase in esculin fluorescence induced by NH 3 indicated some vacuolar alkalization in a Spinacia oleracea L. leaf. Photosynthesis and transpiration remained unchanged during exposure of illuminated leaves to NH 3 for up to 30 min (the maximum tested). CO 2 concentrations influenced the extent of cytosolic alkalinization. 500 μmol CO 2 mol -1 gas suppressed the NH 3-induced cytosolic alkalinization relative to that found in 16 μmol CO 2 mol -1 gas. The suppressing effect of CO 2 on NH 3-induced alkalization was larger in illuminated leaves of the C 4Amaranthus than the C 3Pelargonium. These results indicate that the alkaline pH shift caused by solution and protonation of NH 3 in aqueous leaf compartments is affected by assimilation of NH 3.

C3 and C4 plant responses to increased temperatures and altered monsoonal precipitation in a cool desert on the Colorado Plateau, USA

USGS Publications Warehouse

Wertin, Timothy M.; Reed, Sasha C.; Belnap, Jayne

2015-01-01

Dryland ecosystems represent >40 % of the terrestrial landscape and support over two billion people; consequently, it is vital to understand how drylands will respond to climatic change. However, while arid and semiarid ecosystems commonly experience extremely hot and dry conditions, our understanding of how further temperature increases or altered precipitation will affect dryland plant communities remains poor. To address this question, we assessed plant physiology and growth at a long-term (7-year) climate experiment on the Colorado Plateau, USA, where the community is a mix of shallow-rooted C3 and C4 grasses and deep-rooted C4 shrubs. The experiment maintained elevated-temperature treatments (+2 or +4 °C) in combination with altered summer monsoonal precipitation (+small frequent precipitation events or +large infrequent events). Increased temperature negatively affected photosynthesis and growth of the C3 and C4 grasses, but effects varied in their timing: +4 °C treatments negatively affected the C3 grass early in the growing season of both years, while the negative effects of temperature on the C4 grass were seen in the +2 and +4 °C treatments, but only during the late growing season of the drier year. Increased summer precipitation did not affect photosynthesis or biomass for any species, either in the year the precipitation was applied or the following year. Although previous research suggests dryland plants, and C4 grasses in particular, may respond positively to elevated temperature, our findings from a cool desert show marked declines in C3 and C4 photosynthesis and growth, with temperature effects dependent on the degree of warming and growing-season precipitation.

Hydroxyacetone production from C 3 Criegee intermediates

DOE PAGES

Taatjes, Craig A.; Liu, Fang; Rotavera, Brandon; ...

2016-12-21

Hydroxyacetone (CH 3C(O)CH 2OH) is observed as a stable end product from reactions of the (CH 3) 2COO Criegee intermediate, acetone oxide, in a flow tube coupled with multiplexed photoionization mass spectrometer detection. In the experiment, the isomers at m/z = 74 are distinguished by their different photoionization spectra and reaction times. Hydroxyacetone is observed as a persistent signal at longer reaction times at a higher photoionization threshold of ca. 9.7 eV than Criegee intermediate and definitively identified by comparison with the known photoionization spectrum. Complementary electronic structure calculations reveal multiple possible reaction pathways for hydroxyacetone formation, including unimolecular isomerizationmore » via hydrogen atom transfer and –OH group migration as well as self-reaction of Criegee intermediates. Varying the concentration of Criegee intermediates suggests contributions from both unimolecular and self-reaction pathways to hydroxyacetone. As a result, the hydroxyacetone end product can provide an effective, stable marker for the production of transient Criegee intermediates in future studies of alkene ozonolysis.« less

Phytoremediation of carbamazepine and its metabolite 10,11-epoxycarbamazepine by C3 and C4 plants.

PubMed

Ryšlavá, Helena; Pomeislová, Alice; Pšondrová, Šárka; Hýsková, Veronika; Smrček, Stanislav

2015-12-01

The anticonvulsant drug carbamazepine is considered as an indicator of sewage water pollution: however, its uptake by plants and effect on metabolism have not been sufficiently documented, let alone its metabolite (10,11-epoxycarbamazepine). In a model system of sterile, hydroponically cultivated Zea mays (as C4 plant) and Helianthus annuus (as C3 plant), the uptake and effect of carbamazepine and 10,11-epoxycarbamazepine were studied in comparison with those of acetaminophen and ibuprofen. Ibuprofen and acetaminophen were effectively extracted from drug-supplemented media by both plants, while the uptake of more hydrophobic carbamazepine was much lower. On the other hand, the carbamazepine metabolite, 10,11-epoxycarbamazepine, was, unlike sunflower, willingly taken up by maize plants (after 96 h 88 % of the initial concentration) and effectively stored in maize tissues. In addition, the effect of the studied pharmaceuticals on the plant metabolism (enzymes of Hatch-Slack cycle, peroxidases) was followed. The activity of bound peroxidases, which could cause xylem vessel lignification and reduction of xenobiotic uptake, was at the level of control plants in maize leaves contrary to sunflower. Therefore, our results indicate that maize has the potential to remove 10,11-epoxycarbamazepine from contaminated soils.

Interaction between C 4 barnyard grass and C 3 upland rice under elevated CO 2: Impact of mycorrhizae

NASA Astrophysics Data System (ADS)

Tang, Jianjun; Xu, Liming; Chen, Xin; Hu, Shuijin

2009-03-01

Atmospheric CO 2 enrichment may impact arbuscular mycorrhizae (AM) development and function, which could have subsequent effects on host plant species interactions by differentially affecting plant nutrient acquisition. However, direct evidence illustrating this scenario is limited. We examined how elevated CO 2 affects plant growth and whether mycorrhizae mediate interactions between C 4 barnyard grass ( Echinochloa crusgalli (L.) Beauv.) and C 3 upland rice ( Oryza sativa L.) in a low nutrient soil. The monocultures and combinations with or without mycorrhizal inoculation were grown at ambient (400 ± 20 μmol mol -1) and elevated CO 2 (700 ± 20 μmol mol -1) levels. The 15N isotope tracer was introduced to quantify the mycorrhizally mediated N acquisition of plants. Elevated CO 2 stimulated the growth of C 3 upland rice but not that of C 4 barnyard grass under monoculture. Elevated CO 2 also increased mycorrhizal colonization of C 4 barnyard grass but did not affect mycorrhizal colonization of C 3 upland rice. Mycorrhizal inoculation increased the shoot biomass ratio of C 4 barnyard grass to C 3 upland rice under both CO 2 concentrations but had a greater impact under the elevated than ambient CO 2 level. Mycorrhizae decreased relative interaction index (RII) of C 3 plants under both ambient and elevated CO 2, but mycorrhizae increased RII of C 4 plants only under elevated CO 2. Elevated CO 2 and mycorrhizal inoculation enhanced 15N and total N and P uptake of C 4 barnyard grass in mixture but had no effects on N and P acquisition of C 3 upland rice, thus altering the distribution of N and P between the species in mixture. These results implied that CO 2 stimulation of mycorrhizae and their nutrient acquisition may impact competitive interaction of C 4 barnyard grass and C 3 upland rice under future CO 2 scenarios.

Phenology and productivity of C3 and C4 grasslands in Hawaii.

PubMed

Pau, Stephanie; Still, Christopher J

2014-01-01

Grasslands account for a large proportion of global terrestrial productivity and play a critical role in carbon and water cycling. Within grasslands, photosynthetic pathway is an important functional trait yielding different rates of productivity along environmental gradients. Recently, C3-C4 sorting along spatial environmental gradients has been reassessed by controlling for confounding traits in phylogenetically structured comparisons. C3 and C4 grasses should sort along temporal environmental gradients as well, resulting in differing phenologies and growing season lengths. Here we use 10 years of satellite data (NDVI) to examine the phenology and greenness (as a proxy for productivity) of C3 and C4 grass habitats, which reflect differences in both environment and plant physiology. We perform phylogenetically structured comparisons based on 3,595 digitized herbarium collections of 152 grass species across the Hawaiian Islands. Our results show that the clade identity of grasses captures differences in their habitats better than photosynthetic pathway. Growing season length (GSL) and associated productivity (GSP) were not significantly different when considering photosynthetic type alone, but were indeed different when considering photosynthetic type nested within clade. The relationship between GSL and GSP differed most strongly between C3 clade habitats, and not between C3-C4 habitats. Our results suggest that accounting for the interaction between phylogeny and photosynthetic pathway can help improve predictions of productivity, as commonly used C3-C4 classifications are very broad and appear to mask important diversity in grassland ecosystem functions.

Remotely-sensed phenologies of C3 and C4 grasses in Hawaii using MODIS Vegetation Indices

NASA Astrophysics Data System (ADS)

Pau, S.; Still, C. J.

2010-12-01

The C3 and C4 photosynthetic pathway is a fundamental physiological and ecological distinction in tropical and subtropical savannas and grasslands. Although C4 plants account for 20-25% of global terrestrial productivity, large uncertainties remain regarding their response to climate variability and future climate change. Recent work has shown that key differences in the ecology of C3 and C4 grasses may have been pre-adaptations to environments prior to the evolution of the C4 grasses and not attributable to photosynthetic pathway. The Hawaiian Islands are ideal for studying C3 and C4 grass plant functional types (PFTs) because of the combination of broad climatic gradients within a small geographic area. This study uses MODIS NDVI and EVI time-series data to examine the phenologies of C3 and C4 grasses in a phylogenetic context. Specifically we address 3 primary questions: (1) Do C3 and C4 sister taxa, and C4 subtypes exhibit distinct timing in phenological metrics (onset of greening, onset of senescence, maximum and minimum greenness, length of growing season)? (2) How does the interannual variation in these phenological metrics correlate with interannual variations in climate such as precipitation, air temperature, land surface temperature, and sea surface temperature? (3) How does the length of the growing season translate into differences in productivity?

Drivers of inter-annual variability in C4 abundance in mixed C3-C4 grasslands

NASA Astrophysics Data System (ADS)

Griffith, D.; Ratajczak, Z.; Anderson, M.; Lind, E. M.; Still, C. J.

2016-12-01

Grassland communities tend to be dominated by either C3 or C4 grass species, as opposed to being evenly mixed. Globally, this pattern is a consequence of the crossover temperature threshold above which C4 grasses are climatically favored. However, C3-C4 distributions can also be distinctly bimodal at the landscape scale, reflecting variation in fire regime, herbivory, soils, and other factors that favor either C3 or C4 vegetation. As such, our aims were to first investigate the global controls on C3 and C4 species pools, and second to determine the magnitude of inter-annual variation in C4 grass relative abundance in mixed C3-C4 grasslands with different fire regimes, soil nitrogen, and grazing pressures. Our analyses used data from 74 globally distributed Nutrient Network sites, 30 of which are mixed C3-C4 grasslands. Each site has factorial fertilizer (NPK) and herbivore exclosure treatments in replicated blocks. To address our first goal we conducted a random forest analysis of site-level C4 relative abundances in relation to mean annual temperature and rainfall, growing season temperature (GST) and rainfall, rainfall seasonality, aridity, fire frequency and management, frost, soil fertility, and grass lineage. In order to address our second goal, we narrowed our focus to sites having mixed C3-C4 grass composition and at least five years of species composition data (16 sites). A GST of 15 °C was a good descriptor of C4 versus C3 grass dominance, although there were marked differences among specific C4 grass lineages in their distributions. For example, whether or not a site has an actively managed burn regime was a greater predictor of Andropogoneae (C4) than GST. Furthermore, in mixed C3-C4 grasslands fertilization favored C3 grasses. Our research delineates the climatic limits of mixed C3-C4 grasslands and highlights the influence of disturbance, soil, and phylogeny on C4 and C3 grass dominance.

Comparative studies of C3 and C4 Atriplex hybrids in the genomics era: physiological assessments

PubMed Central

Oakley, Jason C.; Sultmanis, Stefanie; Stinson, Corey R.; Sage, Tammy L.; Sage, Rowan F.

2014-01-01

We crossed the C3 species Atriplex prostrata with the C4 species Atriplex rosea to produce F1 and F2 hybrids. All hybrids exhibited C3-like δ13C values, and had reduced rates of net CO2 assimilation compared with A. prostrata. The activities of the major C4 cycle enzymes PEP carboxylase, NAD-malic enzyme, and pyruvate-Pi dikinase in the hybrids were at most 36% of the C4 values. These results demonstrate the C4 metabolic cycle was disrupted in the hybrids. Photosynthetic CO2 compensation points (Г) of the hybrids were generally midway between the C3 and C4 values, and in most hybrids were accompanied by low, C3-like activities in one or more of the major C4 cycle enzymes. This supports the possibility that most hybrids use a photorespiratory glycine shuttle to concentrate CO2 into the bundle sheath cells. One hybrid exhibited a C4-like Г of 4 µmol mol–1, indicating engagement of a C4 metabolic cycle. Consistently, this hybrid had elevated activities of all measured C4 cycle enzymes relative to the C3 parent; however, C3-like carbon isotope ratios indicate the low Г is mainly due to a photorespiratory glycine shuttle. The anatomy of the hybrids resembled that of C3-C4 intermediate species using a glycine shuttle to concentrate CO2 in the bundle sheath, and is further evidence that this physiology is the predominant, default condition of the F2 hybrids. Progeny of these hybrids should further segregate C3 and C4 traits and in doing so assist in the discovery of C4 genes using high-throughput methods of the genomics era. PMID:24675672

The Paleo-ecology of C4 Evolution

NASA Astrophysics Data System (ADS)

Sage, R. F.; Khoshravesh, R.

2014-12-01

Molecular clock analysis of extant plant lineages consistently place the earliest appearance of the C4 photosynthetic pathway in the mid-to-late Oligocene, coincident with a decline in atmospheric CO2 and a spread of dry environments. Most of the approximately 70 known lineages of C4 photosynthesis, however, evolved over the subsequent 23 million years since the Oligocene. Examination of living C3-C4 intermediate species, and close C3 relatives of modern C4 lineages, indicate that the C4 pathway evolved in regions of high heat and episodic drought and/or salinity, usually in the drier ends of the monsoon belts of the subtropics. Soils associated with transitional species are typically sandy, rocky, or salinized, and have low vegetation density, which in combination with high air temperature allows for high surface heat loads that warm leaves to near 45°C. Under such conditions in low CO2 atmospheres, the rate of photorespiration is very high and would greatly impair C3 photosynthesis and establish conditions favoring C4 evolution. However, studies with modern taxa do not address whether the extreme habitats proposed to facilitate C4 evolution were actually present at the time when the C4 pathway evolved in any given lineage. Here, we examine the paleo-record to evaluate the environmental conditions present in the C4 centres of origin when the respective transitions from C3 to C4 photosynthesis are estimated to have occurred.

Reduced plant water status under sub-ambient pCO2 limits plant productivity in the wild progenitors of C3 and C4 cereals

PubMed Central

Cunniff, Jennifer; Charles, Michael; Jones, Glynis; Osborne, Colin P.

2016-01-01

Background and Aims The reduction of plant productivity by low atmospheric CO2 partial pressure (pCO2) during the last glacial period is proposed as a limiting factor for the establishment of agriculture. Supporting this hypothesis, previous work has shown that glacial pCO2 limits biomass in the wild progenitors of C3 and C4 founder crops, in part due to the direct effects of glacial pCO2 on photosynthesis. Here, we investigate the indirect role of pCO2 mediated via water status, hypothesizing that faster soil water depletion at glacial (18 Pa) compared to post-glacial (27 Pa) pCO2, due to greater stomatal conductance, feeds back to limit photosynthesis during drying cycles. Methods We grew four wild progenitors of C3 and C4 crops at glacial and post-glacial pCO2 and investigated physiological changes in gas exchange, canopy transpiration, soil water content and water potential between regular watering events. Growth parameters including leaf area were measured. Key Results Initial transpiration rates were higher at glacial pCO2 due to greater stomatal conductance. However, stomatal conductance declined more rapidly over the soil drying cycle in glacial pCO2 and was associated with decreased intercellular pCO2 and lower photosynthesis. Soil water content was similar between pCO2 levels as larger leaf areas at post-glacial pCO2 offset the slower depletion of water. Instead the feedback could be linked to reduced plant water status. Particularly in the C4 plants, soil–leaf water potential gradients were greater at 18 Pa compared with 27 Pa pCO2, suggesting an increased ratio of leaf evaporative demand to supply. Conclusions Reduced plant water status appeared to cause a negative feedback on stomatal aperture in plants at glacial pCO2, thereby reducing photosynthesis. The effects were stronger in C4 species, providing a mechanism for reduced biomass at 18 Pa. These results have added significance when set against the drier climate of the glacial period

Glyphosate Resistance of C3 and C4 Weeds under Rising Atmospheric CO2.

PubMed

Fernando, Nimesha; Manalil, Sudheesh; Florentine, Singarayer K; Chauhan, Bhagirath S; Seneweera, Saman

2016-01-01

The present paper reviews current knowledge on how changes of plant metabolism under elevated CO2 concentrations (e[CO2]) can affect the development of the glyphosate resistance of C3 and C4 weeds. Among the chemical herbicides, glyphosate, which is a non-selective and post-emergence herbicide, is currently the most widely used herbicide in global agriculture. As a consequence, glyphosate resistant weeds, particularly in major field crops, are a widespread problem and are becoming a significant challenge to future global food production. Of particular interest here it is known that the biochemical processes involved in photosynthetic pathways of C3 and C4 plants are different, which may have relevance to their competitive development under changing environmental conditions. It has already been shown that plant anatomical, morphological, and physiological changes under e[CO2] can be different, based on (i) the plant's functional group, (ii) the available soil nutrients, and (iii) the governing water status. In this respect, C3 species are likely to have a major developmental advantage under a CO2 rich atmosphere, by being able to capitalize on the overall stimulatory effect of e[CO2]. For example, many tropical weed grass species fix CO2 from the atmosphere via the C4 photosynthetic pathway, which is a complex anatomical and biochemical variant of the C3 pathway. Thus, based on our current knowledge of CO2 fixing, it would appear obvious that the development of a glyphosate-resistant mechanism would be easier under an e[CO2] in C3 weeds which have a simpler photosynthetic pathway, than for C4 weeds. However, notwithstanding this logical argument, a better understanding of the biochemical, genetic, and molecular measures by which plants develop glyphosate resistance and how e[CO2] affects these measures will be important before attempting to innovate sustainable technology to manage the glyphosate-resistant evolution of weeds under e[CO2]. Such information will be of

The Photo-3 model: A Python-based model for C3, C4, and CAM photosynthesis coupled with environmental conditions

NASA Astrophysics Data System (ADS)

Hartzell, S. R.; Bartlett, M. S., Jr.; Porporato, A. M.

2017-12-01

The ability to depict all three photosynthetic types (C3, C4, and CAM) has important implications for the study of both natural and agroecosystems. Currently no model exists which covers all types of photosynthesis in a consistent way and which can be fully integrated with environmental conditions. This is partially because, despite the fact that Crassulacean acid metabolism (CAM) photosynthesis is prevalent in many plants in arid and semi-arid ecosystems, where it may comprise nearly 50% of all plant biomass, CAM modelling remains understudied. The Photo-3 model takes advantage of recent advances in mechanistic modeling of CAM photosynthesis to provide a direct comparison of CAM functioning with C3 and C4 functioning under a wide range of soil and atmospheric conditions. The model is based on a core Farquhar photosynthetic model with additional functions to represent the spatial and temporal separations of carbon uptake and assimilation in the case of C4 and CAM photosynthesis. We have parameterized the model for one representative species of each photosynthetic type: Opuntia ficus-indica (CAM), Sorghum bicolor (C4), and Triticum aestivum (C3). Results agree well with experimental data on carbon assimilation and water use for the three species. Model runs using climate data from Temple, TX; Sicily, Italy; Zacatecas, Mexico; Pernambuco, Brazil and Adias Ababa, Ethiopia illustrate the high water use efficiency of CAM plants and its cumulative effects on long-term productivity in water-limited environments. The Photo-3 model, which is written in Python, will be made publicly available on GitHub and its outputs may be coupled to existing models of plant growth and phenology. The model may be used to evaluate potential productivity and water use for C3, C4, and CAM plants, and to devise optimal strategies for cropping systems and irrigation in water-limited environments.

Germination sensitivities to water potential among co-existing C3 and C4 grasses of cool semi-arid prairie grasslands.

PubMed

Mollard, F P O; Naeth, M A

2015-03-01

An untested theory states that C4 grass seeds could germinate under lower water potentials (Ψ) than C3 grass seeds. We used hydrotime modelling to study seed water relations of C4 and C3 Canadian prairie grasses to address Ψ divergent sensitivities and germination strategies along a risk-spreading continuum of responses to limited water. C4 grasses were Bouteloua gracilis, Calamovilfa longifolia and Schizachyrium scoparium; C3 grasses were Bromus carinatus, Elymus trachycaulus, Festuca hallii and Koeleria macrantha. Hydrotime parameters were obtained after incubation of non-dormant seeds under different Ψ PEG 6000 solutions. A t-test between C3 and C4 grasses did not find statistical differences in population mean base Ψ (Ψb (50)). We found idiosyncratic responses of C4 grasses along the risk-spreading continuum. B. gracilis showed a risk-taker strategy of a species able to quickly germinate in a dry soil due to its low Ψb (50) and hydrotime (θH ). The high Ψb (50) of S. scoparium indicates it follows the risk-averse strategy so it can only germinate in wet soils. C. longifolia showed an intermediate strategy: the lowest Ψb (50) yet the highest θH . K. macrantha, a C3 grass which thrives in dry habitats, had the highest Ψb (50), suggesting a risk-averse strategy for a C3 species. Other C3 species showed intermediate germination patterns in response to Ψ relative to C4 species. Our results indicate that grasses display germination sensitivities to Ψ across the risk-spreading continuum of responses. Thus seed water relations may be poor predictors to explain differential recruitment and distribution of C3 and C4 grasses in the Canadian prairies. © 2014 German Botanical Society and The Royal Botanical Society of the Netherlands.

Germination shifts of C3 and C4 species under simulated global warming scenario.

PubMed

Zhang, Hongxiang; Yu, Qiang; Huang, Yingxin; Zheng, Wei; Tian, Yu; Song, Yantao; Li, Guangdi; Zhou, Daowei

2014-01-01

Research efforts around the world have been increasingly devoted to investigating changes in C3 and C4 species' abundance or distribution with global warming, as they provide important insight into carbon fluxes and linked biogeochemical cycles. However, changes in the early life stage (e.g. germination) of C3 and C4 species in response to global warming, particularly with respect to asymmetric warming, have received less attention. We investigated germination percentage and rate of C3 and C4 species under asymmetric (+3/+6°C at day/night) and symmetric warming (+5/+5°C at day/night), simulated by alternating temperatures. A thermal time model was used to calculate germination base temperature and thermal time constant. Two additional alternating temperature regimes were used to test temperature metrics effect. The germination percentage and rate increased continuously for C4 species, but increased and then decreased with temperature for C3 species under both symmetric and asymmetric warming. Compared to asymmetric warming, symmetric warming significantly overestimated the speed of germination percentage change with temperature for C4 species. Among the temperature metrics (minimum, maximum, diurnal temperature range and average temperature), maximum temperature was most correlated with germination of C4 species. Our results indicate that global warming may favour germination of C4 species, at least for the C4 species studied in this work. The divergent effects of asymmetric and symmetric warming on plant germination also deserve more attention in future studies.

Glyphosate Resistance of C3 and C4 Weeds under Rising Atmospheric CO2

PubMed Central

Fernando, Nimesha; Manalil, Sudheesh; Florentine, Singarayer K.; Chauhan, Bhagirath S.; Seneweera, Saman

2016-01-01

The present paper reviews current knowledge on how changes of plant metabolism under elevated CO2 concentrations (e[CO2]) can affect the development of the glyphosate resistance of C3 and C4 weeds. Among the chemical herbicides, glyphosate, which is a non-selective and post-emergence herbicide, is currently the most widely used herbicide in global agriculture. As a consequence, glyphosate resistant weeds, particularly in major field crops, are a widespread problem and are becoming a significant challenge to future global food production. Of particular interest here it is known that the biochemical processes involved in photosynthetic pathways of C3 and C4 plants are different, which may have relevance to their competitive development under changing environmental conditions. It has already been shown that plant anatomical, morphological, and physiological changes under e[CO2] can be different, based on (i) the plant’s functional group, (ii) the available soil nutrients, and (iii) the governing water status. In this respect, C3 species are likely to have a major developmental advantage under a CO2 rich atmosphere, by being able to capitalize on the overall stimulatory effect of e[CO2]. For example, many tropical weed grass species fix CO2 from the atmosphere via the C4 photosynthetic pathway, which is a complex anatomical and biochemical variant of the C3 pathway. Thus, based on our current knowledge of CO2 fixing, it would appear obvious that the development of a glyphosate-resistant mechanism would be easier under an e[CO2] in C3 weeds which have a simpler photosynthetic pathway, than for C4 weeds. However, notwithstanding this logical argument, a better understanding of the biochemical, genetic, and molecular measures by which plants develop glyphosate resistance and how e[CO2] affects these measures will be important before attempting to innovate sustainable technology to manage the glyphosate-resistant evolution of weeds under e[CO2]. Such information will be

C3 and C4 biomass allocation responses to elevated CO2 and nitrogen: contrasting resource capture strategies

USGS Publications Warehouse

White, K.P.; Langley, J.A.; Cahoon, D.R.; Megonigal, J.P.

2012-01-01

Plants alter biomass allocation to optimize resource capture. Plant strategy for resource capture may have important implications in intertidal marshes, where soil nitrogen (N) levels and atmospheric carbon dioxide (CO2) are changing. We conducted a factorial manipulation of atmospheric CO2 (ambient and ambient + 340 ppm) and soil N (ambient and ambient + 25 g m-2 year-1) in an intertidal marsh composed of common North Atlantic C3 and C4 species. Estimation of C3 stem turnover was used to adjust aboveground C3 productivity, and fine root productivity was partitioned into C3-C4 functional groups by isotopic analysis. The results suggest that the plants follow resource capture theory. The C3 species increased aboveground productivity under the added N and elevated CO2 treatment (P 2 alone. C3 fine root production decreased with added N (P 2 (P = 0.0481). The C4 species increased growth under high N availability both above- and belowground, but that stimulation was diminished under elevated CO2. The results suggest that the marsh vegetation allocates biomass according to resource capture at the individual plant level rather than for optimal ecosystem viability in regards to biomass influence over the processes that maintain soil surface elevation in equilibrium with sea level.

On the contributions of photorespiration and compartmentation to the contrasting intramolecular 2H profiles of C3 and C4 plant sugars

Treesearch

Youping Zhou; Benli Zhang; Hilary Stuart-Williams; Kliti Grice; Charles H. Hocart; Arthur Gessler; Zachary E. Kayler; Graham D. Farquhar

2018-01-01

Compartmentation of C4 photosynthetic biochemistry into bundle sheath (BS) and mesophyll (M) cells, and photorespiration in C3 plants is predicted to have hydrogen isotopic consequences for metabolites at both molecular and site-specific levels. Molecular-level evidence was recently reported (Zhou et al., 2016), but...

Temporal and spatial variations of canopy temperature over a C3C4 mixture grassland

NASA Astrophysics Data System (ADS)

Shimoda, S.; Oikawa, T.

2006-10-01

This study discusses the photosynthetic pathway types involved in canopy temperature measurements on a mixed grassland consisting of C3 and C4 plants (dominant species in biomass were Solidago altissima (C3), Miscanthus sinensis (C4), and Imperata cylindrica (C4)). In the wet conditions immediately after the rainy season, the mean canopy temperature for S. altissima was the lowest among the dominant species, mainly due to its leaf conductance being twice as large as the other two species. Despite using the same C4 photosynthetic pathway, M. sinensis had a lower apparent canopy temperature than I. cylindrica due to a smaller proportion of sunlit elements in the field of view. In the dry conditions during late July, the mean canopy temperatures of the three dominant species were within 0.3 °C of one another. These results can be explained by poor water conditions for C3 species (S. altissima). The simultaneous survey of vegetation and thermal imaging can help clarify characteristics of C3 and C4 canopy temperature over complicated grassland.

Walking the C4 pathway: past, present, and future.

PubMed

Furbank, Robert T

2017-01-01

The year 2016 marks 50 years since the publication of the seminal paper by Hatch and Slack describing the biochemical pathway we now know as C 4 photosynthesis. This review provides insight into the initial discovery of this pathway, the clues which led Hatch and Slack and others to these definitive experiments, some of the intrigue which surrounds the international activities which led up to the discovery, and personal insights into the future of this research field. While the biochemical understanding of the basic pathways came quickly, the role of the bundle sheath intermediate CO 2 pool was not understood for a number of years, and the nature of C 4 as a biochemical CO 2 pump then linked the unique Kranz anatomy of C 4 plants to their biochemical specialization. Decades of "grind and find biochemistry" and leaf physiology fleshed out the regulation of the pathway and the differences in physiological response to the environment between C 3 and C 4 plants. The more recent advent of plant transformation then high-throughput RNA and DNA sequencing and synthetic biology has allowed us both to carry out biochemical experiments and test hypotheses in planta and to better understand the evolution-driven molecular and genetic changes which occurred in the genomes of plants in the transition from C 3 to C 4 Now we are using this knowledge in attempts to engineer C 4 rice and improve the C 4 engine itself for enhanced food security and to provide novel biofuel feedstocks. The next 50 years of photosynthesis will no doubt be challenging, stimulating, and a drawcard for the best young minds in plant biology. © The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.

Walking the C4 pathway: past, present, and future.

PubMed

Furbank, Robert T

2016-07-01

The year 2016 marks 50 years since the publication of the seminal paper by Hatch and Slack describing the biochemical pathway we now know as C4 photosynthesis. This review provides insight into the initial discovery of this pathway, the clues which led Hatch and Slack and others to these definitive experiments, some of the intrigue which surrounds the international activities which led up to the discovery, and personal insights into the future of this research field. While the biochemical understanding of the basic pathways came quickly, the role of the bundle sheath intermediate CO2 pool was not understood for a number of years, and the nature of C4 as a biochemical CO2 pump then linked the unique Kranz anatomy of C4 plants to their biochemical specialization. Decades of "grind and find biochemistry" and leaf physiology fleshed out the regulation of the pathway and the differences in physiological response to the environment between C3 and C4 plants. The more recent advent of plant transformation then high-throughput RNA and DNA sequencing and synthetic biology has allowed us both to carry out biochemical experiments and test hypotheses in planta and to better understand the evolution-driven molecular and genetic changes which occurred in the genomes of plants in the transition from C3 to C4 Now we are using this knowledge in attempts to engineer C4 rice and improve the C4 engine itself for enhanced food security and to provide novel biofuel feedstocks. The next 50 years of photosynthesis will no doubt be challenging, stimulating, and a drawcard for the best young minds in plant biology. © The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.

Reactive oxygen species and redox regulation in mesophyll and bundle sheath cells of C4 plants.

PubMed

Turkan, Ismail; Uzilday, Baris; Dietz, Karl-Josef; Bräutigam, Andrea; Ozgur, Rengin

2018-02-26

Redox regulation, antioxidant defence and ROS signalling are critical in realizing and tuning metabolic activities. However, our concepts were mostly developed for C3 plants since Arabidopsis thaliana is major model. Efforts to convert C3 plants to C4 plants to increase yield (see C4 rice; c4rice.irri.org/) entails better understanding of these processes in C4 plants. Various photosynthetic enzymes that take part in light reactions and carbon reactions are regulated via redox components such as thioredoxins as redox transmitters and peroxiredoxins. Due to this, understanding redox regulation in mesophyll and bundle sheath chloroplasts of C4 plants is of paramount importance. It appears impossible to utilize efficient C4 photosynthesis without understanding its exact redox needs and regulation mechanisms used during light reactions. In this review we will discuss available knowledge on redox regulation in C3 and C4 plants with special emphasis on mesophyll and bundle sheath differences in C4. In these two cell types of C4 plants, linear and cyclic electron transport in chloroplasts operate differentially when compared to C3 chloroplasts, changing the redox needs of the cell. Therefore, the focus is given to photosynthetic light reactions, ROS production dynamics, antioxidant defence and thiol based redox regulation with the aim to draw a picture of current knowledge.

Effects of Botrytis cinerea and Pseudomonas syringae infection on the antioxidant profile of Mesembryanthemum crystallinum C3/CAM intermediate plant.

PubMed

Libik-Konieczny, Marta; Surówka, Ewa; Kuźniak, Elżbieta; Nosek, Michał; Miszalski, Zbigniew

2011-07-01

Mesembryathemum crystallinum plants performing C(3) or CAM (crassulacean acid metabolism) appear to be highly resistant to Botrytis cinerea as well as to Pseudomonas syringae. Fungal hyphae growth was restricted to 48h post-inoculation (hpi) in both metabolic types and morphology of hyphae differed between those growing in C(3) and CAM plants. Growth of bacteria was inhibited significantly 24 hpi in both C(3) and CAM plants. B. cinerea and P. syringae infection led to an increase in the concentration of H(2)O(2) in C(3) plants 3 hpi, while a decrease in H(2)O(2) content was observed in CAM performing plants. The concentration of H(2)O(2) returned to the control level 24 and 48 hpi. Changes in H(2)O(2) content corresponded with the activity of guaiacol peroxidase (POD), mostly 3 hpi. We noted that its activity decreased significantly in C(3) plants and increased in CAM plants in response to inoculation with both pathogens. On the contrary, changes in the activity of CAT did not correlate with H(2)O(2) level. It increased significantly after interaction of C(3) plants with B. cinerea or P. syringae, but in CAM performing plants, the activity of this enzyme was unchanged. Inoculation with B. cinerea or P. syringae led to an increase in the total SOD activity in C(3) plants while CAM plants did not exhibit changes in the total SOD activity after interaction with both pathogens. In conclusion, the pathogen-induced changes in H(2)O(2) content and in SOD, POD and CAT activities in M. crystallinum leaves, were related to the photosynthetic metabolism type of the stressed plants rather than to the lifestyle of the invading pathogen. Copyright © 2011 Elsevier GmbH. All rights reserved.

Carbon isotopic evidence from paleosols for mixed C 3/C 4 vegetation in the Bogota Basin, Colombia

NASA Astrophysics Data System (ADS)

Mora, Germán; Pratt, Lisa M.

2002-04-01

Pollen reconstructions in the Bogota basin (Colombia) indicate the expansion of tropical high-altitude grassland (paramo) at the expense of Andean forests during glacial intervals. The carbon isotopic composition (δ 13C) of soil organic matter (SOM) can be a useful indicator of changes in vegetation affecting grasslands because it distinguishes between two groups of grasses (C 3 and C 4) adapted to different ecological environments. Values of SOM δ 13C were determined in four weathering profiles containing both modern (Holocene) soils and paleosols formed during the Last Glacial Stage. These profiles are located along an altitudinal transect in the Bogota basin, extending from 2550 to 3100 m. Values of SOM δ 13C in the topsoil horizons reflect those of the native C 3 vegetation that currently dominates the ecosystems in the Colombian Andes. Although C 4 grasses are currently negligible in the basin, elevated SOM δ 13C values indicative of C 4 plants were found in two Holocene soils. Environmental changes or ancient agricultural activities could explain the increased abundance of these plants in the basin during the late Holocene. Isotopic values in the studied paleosols revealed the presence of a mixed C 3/C 4 vegetation in the basin during the Last Glacial Stage, thus indicating the expansion of C 4 grasses. We hypothesized that lowered pCO 2 and possibly reduced rainfall resulted in the colonization of the tropical Andes by lowland C 4 grasses despite of prevailing cooler temperatures.

Improved analysis of C4 and C3 photosynthesis via refined in vitro assays of their carbon fixation biochemistry

PubMed Central

Sharwood, Robert E.; Sonawane, Balasaheb V.; Ghannoum, Oula; Whitney, Spencer M.

2016-01-01

Plants operating C3 and C4 photosynthetic pathways exhibit differences in leaf anatomy and photosynthetic carbon fixation biochemistry. Fully understanding this underpinning biochemical variation is requisite to identifying solutions for improving photosynthetic efficiency and growth. Here we refine assay methods for accurately measuring the carboxylase and decarboxylase activities in C3 and C4 plant soluble protein. We show that differences in plant extract preparation and assay conditions are required to measure NADP-malic enzyme and phosphoenolpyruvate carboxylase (pH 8, Mg2+, 22 °C) and phosphoenolpyruvate carboxykinase (pH 7, >2mM Mn2+, no Mg2+) maximal activities accurately. We validate how the omission of MgCl2 during leaf protein extraction, lengthy (>1min) centrifugation times, and the use of non-pure ribulose-1,5-bisphosphate (RuBP) significantly underestimate Rubisco activation status. We show how Rubisco activation status varies with leaf ontogeny and is generally lower in mature C4 monocot leaves (45–60% activation) relative to C3 monocots (55–90% activation). Consistent with their >3-fold lower Rubisco contents, full Rubisco activation in soluble protein from C4 leaves (<5min) was faster than in C3 plant samples (<10min), with addition of Rubisco activase not required for full activation. We conclude that Rubisco inactivation in illuminated leaves primarily stems from RuBP binding to non-carbamylated enzyme, a state readily reversible by dilution during cellular protein extraction. PMID:27122573

Chalcone-based Selective Inhibitors of a C4 Plant Key Enzyme as Novel Potential Herbicides

NASA Astrophysics Data System (ADS)

Nguyen, G. T. T.; Erlenkamp, G.; Jäck, O.; Küberl, A.; Bott, M.; Fiorani, F.; Gohlke, H.; Groth, G.

2016-06-01

Weeds are a challenge for global food production due to their rapidly evolving resistance against herbicides. We have identified chalcones as selective inhibitors of phosphoenolpyruvate carboxylase (PEPC), a key enzyme for carbon fixation and biomass increase in the C4 photosynthetic pathway of many of the world’s most damaging weeds. In contrast, many of the most important crop plants use C3 photosynthesis. Here, we show that 2‧,3‧,4‧,3,4-Pentahydroxychalcone (IC50 = 600 nM) and 2‧,3‧,4‧-Trihydroxychalcone (IC50 = 4.2 μM) are potent inhibitors of C4 PEPC but do not affect C3 PEPC at a same concentration range (selectivity factor: 15-45). Binding and modeling studies indicate that the active compounds bind at the same site as malate/aspartate, the natural feedback inhibitors of the C4 pathway. At the whole plant level, both substances showed pronounced growth-inhibitory effects on the C4 weed Amaranthus retroflexus, while there were no measurable effects on oilseed rape, a C3 plant. Growth of selected soil bacteria was not affected by these substances. Our chalcone compounds are the most potent and selective C4 PEPC inhibitors known to date. They offer a novel approach to combat C4 weeds based on a hitherto unexplored mode of allosteric inhibition of a C4 plant key enzyme.

Chalcone-based Selective Inhibitors of a C4 Plant Key Enzyme as Novel Potential Herbicides

PubMed Central

Nguyen, G. T. T.; Erlenkamp, G.; Jäck, O.; Küberl, A.; Bott, M.; Fiorani, F.; Gohlke, H.; Groth, G.

2016-01-01

Weeds are a challenge for global food production due to their rapidly evolving resistance against herbicides. We have identified chalcones as selective inhibitors of phosphoenolpyruvate carboxylase (PEPC), a key enzyme for carbon fixation and biomass increase in the C4 photosynthetic pathway of many of the world’s most damaging weeds. In contrast, many of the most important crop plants use C3 photosynthesis. Here, we show that 2′,3′,4′,3,4-Pentahydroxychalcone (IC50 = 600 nM) and 2′,3′,4′-Trihydroxychalcone (IC50 = 4.2 μM) are potent inhibitors of C4 PEPC but do not affect C3 PEPC at a same concentration range (selectivity factor: 15–45). Binding and modeling studies indicate that the active compounds bind at the same site as malate/aspartate, the natural feedback inhibitors of the C4 pathway. At the whole plant level, both substances showed pronounced growth-inhibitory effects on the C4 weed Amaranthus retroflexus, while there were no measurable effects on oilseed rape, a C3 plant. Growth of selected soil bacteria was not affected by these substances. Our chalcone compounds are the most potent and selective C4 PEPC inhibitors known to date. They offer a novel approach to combat C4 weeds based on a hitherto unexplored mode of allosteric inhibition of a C4 plant key enzyme. PMID:27263468

Africa's wild C4 plant foods and possible early hominid diets.

PubMed

Peters, Charles R; Vogel, John C

2005-03-01

A small minority of Africa's wild plant foods are C4. These are primarily the seeds of some of the C4 grasses, the rootstocks and stem/leaf bases of some of the C4 sedges (especially papyrus), and the leaves of some of the C4 herbaceous dicots (forbs). These wild food plants are commonly found in disturbed ground and wetlands (particularly the grasses and sedges). Multiple lines of evidence indicate that C4 grasses were present in Africa by at least the late Miocene. It is a reasonable hypothesis that the prehistory of the C4 sedges parallels that of the C4 grasses, but the C4 forbs may not have become common until the late Pleistocene. CAM plants may have a more ancient history, but offer few opportunities for an additional C4-like dietary signal. The environmental reconstructions available for the early South African hominid sites do not indicate the presence of large wetlands, and therefore probably the absence of a strong potential for a C4 plant food diet. However, carbon isotope analyses of tooth enamel from three species of early South African hominids have shown that there was a significant but not dominant contribution of C4 biomass in their diets. Since it appears unlikely that this C4 component could have come predominantly from C4 plant foods, a broad range of potential animal contributors is briefly considered, namely invertebrates, reptiles, birds, and small mammals. It is concluded that the similar average C4 dietary intake seen in the three South African hominid species could have been acquired by differing contributions from the various sources, without the need to assume scavenging or hunting of medium to large grazing ungulates. Effectively similar dominantly dryland paleo-environments may also be part of the explanation. Theoretically, elsewhere in southern and eastern Africa, large wetlands would have offered early hominids greater opportunities for a C4 plant diet.

Phenotypic landscape inference reveals multiple evolutionary paths to C4 photosynthesis

PubMed Central

Williams, Ben P; Johnston, Iain G; Covshoff, Sarah; Hibberd, Julian M

2013-01-01

C4 photosynthesis has independently evolved from the ancestral C3 pathway in at least 60 plant lineages, but, as with other complex traits, how it evolved is unclear. Here we show that the polyphyletic appearance of C4 photosynthesis is associated with diverse and flexible evolutionary paths that group into four major trajectories. We conducted a meta-analysis of 18 lineages containing species that use C3, C4, or intermediate C3–C4 forms of photosynthesis to parameterise a 16-dimensional phenotypic landscape. We then developed and experimentally verified a novel Bayesian approach based on a hidden Markov model that predicts how the C4 phenotype evolved. The alternative evolutionary histories underlying the appearance of C4 photosynthesis were determined by ancestral lineage and initial phenotypic alterations unrelated to photosynthesis. We conclude that the order of C4 trait acquisition is flexible and driven by non-photosynthetic drivers. This flexibility will have facilitated the convergent evolution of this complex trait. DOI: http://dx.doi.org/10.7554/eLife.00961.001 PMID:24082995

The soil microbial community composition and soil microbial carbon uptake are more affected by soil type than by different vegetation types (C3 and C4 plants) and seasonal changes

NASA Astrophysics Data System (ADS)

Griselle Mellado Vazquez, Perla; Lange, Markus; Gleixner, Gerd

2016-04-01

This study investigates the influence of different vegetation types (C3 and C4 plants), soil type and seasonal changes on the soil microbial biomass, soil microbial community composition and soil microbial carbon (C) uptake. We collected soil samples in winter (non-growing season) and summer (growing season) in 2012 from an experimental site cropping C3 and C4 plants for 6 years on two different soil types (sandy and clayey). The amount of phospholipid fatty acids (PLFAs) and their compound-specific δ13C values were used to determined microbial biomass and the flow of C from plants to soil microorganisms, respectively. Higher microbial biomass was found in the growing season. The microbial community composition was mainly explained by soil type. Higher amounts of SOC were driving the predominance of G+ bacteria, actinobacteria and cyclic G- bacteria in sandy soils, whereas root biomass was significantly related to the increased proportions of G- bacteria in clayey soils. Plant-derived C in G- bacteria increased significantly in clayey soils in the growing season. This increase was positively and significantly driven by root biomass. Moreover, changes in plant-derived C among microbial groups pointed to specific capabilities of different microbial groups to decompose distinct sources of C. We concluded that soil texture and favorable growth conditions driven by rhizosphere interactions are the most important factors controlling the soil microbial community. Our results demonstrate that a change of C3 plants vs. C4 plants has only a minor effect on the soil microbial community. Thus, such experiments are well suited to investigate soil organic matter dynamics as they allow to trace the C flow from plants into the soil microbial community without changing the community abundance and composition.

Loss of the Chloroplast Transit Peptide from an Ancestral C3 Carbonic Anhydrase Is Associated with C4 Evolution in the Grass Genus Neurachne.

PubMed

Clayton, Harmony; Saladié, Montserrat; Rolland, Vivien; Sharwood, Robert; Macfarlane, Terry; Ludwig, Martha

2017-03-01

Neurachne is the only known grass lineage containing closely related C 3 , C 3 -C 4 intermediate, and C 4 species, making it an ideal taxon with which to study the evolution of C 4 photosynthesis in the grasses. To begin dissecting the molecular changes that led to the evolution of C 4 photosynthesis in this group, the complementary DNAs encoding four distinct β-carbonic anhydrase (CA) isoforms were characterized from leaf tissue of Neurachne munroi (C 4 ), Neurachne minor (C 3 -C 4 ), and Neurachne alopecuroidea (C 3 ). Two genes ( CA1 and CA2 ) each encode two different isoforms: CA1a/CA1b and CA2a/CA2b. Transcript analyses found that CA1 messenger RNAs were significantly more abundant than transcripts from the CA2 gene in the leaves of each species examined, constituting ∼99% of all β-CA transcripts measured. Localization experiments using green fluorescent protein fusion constructs showed that, while CA1b is a cytosolic CA in all three species, the CA1a proteins are differentially localized. The N. alopecuroidea and N. minor CA1a isoforms were imported into chloroplasts of Nicotiana benthamiana leaf cells, whereas N. munroi CA1a localized to the cytosol. Sequence analysis indicated an 11-amino acid deletion in the amino terminus of N. munroi CA1a relative to the C 3 and C 3 -C 4 proteins, suggesting that chloroplast targeting of CA1a is the ancestral state and that loss of a functional chloroplast transit peptide in N. munroi CA1a is associated with the evolution of C 4 photosynthesis in Neurachne spp. Remarkably, this mechanism is homoplastic with the evolution of the C 4 -associated CA in the dicotyledonous genus Flaveria , although the actual mutations in the two lineages differ. © 2017 American Society of Plant Biologists. All Rights Reserved.

Photosynthetic responses of C3 and C4 species to seasonal water variability and competition.

PubMed

Niu, Shuli; Yuan, Zhiyou; Zhang, Yanfang; Liu, Weixing; Zhang, Lei; Huang, Jianhui; Wan, Shiqiang

2005-11-01

This study examined the impacts of seasonal water variability and interspecific competition on the photosynthetic characteristics of a C3 (Leymus chinensis) and a C4 (Chloris virgata) grass species. Plants received the same amount of water but in three seasonal patterns, i.e. the one-peak model (more water in the summer than in the spring and autumn), the two-peak model (more water in the spring and autumn than in the summer), and the average model (water evenly distributed over the growing season). The effects of water variability on the photosynthetic characteristics of the C3 and C4 species were dependent on season. There were significant differences in the photosynthetic characteristics of the C4 species in the summer and the C3 species in the autumn among the three water treatments. Interspecific competition exerted negative impacts on the C3 species in August and September but had no effects on the C4 species in any of the four measuring dates. The relative competitive capability of the two species was not altered by water availability. The assimilation rate, the maximum quantum yield of net CO2 assimilation, and the maximum rate of carboxylation of the C3 species were 13-56%, 5-11%, and 11-48% greater, respectively, in a monoculture than in a mixture in August and September. The results demonstrated that the photosynthetic characteristics of the C3 and C4 species were affected by water availability, but the effects varied considerably with season.

C4 Cycles: Past, Present, and Future Research on C4 Photosynthesis

PubMed Central

Langdale, Jane A.

2011-01-01

In the late 1960s, a vibrant new research field was ignited by the discovery that instead of fixing CO2 into a C3 compound, some plants initially fix CO2 into a four-carbon (C4) compound. The term C4 photosynthesis was born. In the 20 years that followed, physiologists, biochemists, and molecular and developmental biologists grappled to understand how the C4 photosynthetic pathway was partitioned between two morphologically distinct cell types in the leaf. By the early 1990s, much was known about C4 biochemistry, the types of leaf anatomy that facilitated the pathway, and the patterns of gene expression that underpinned the biochemistry. However, virtually nothing was known about how the pathway was regulated. It should have been an exciting time, but many of the original researchers were approaching retirement, C4 plants were proving recalcitrant to genetic manipulation, and whole-genome sequences were not even a dream. In combination, these factors led to reduced funding and the failure to attract young people into the field; the endgame seemed to be underway. But over the last 5 years, there has been a resurgence of interest and funding, not least because of ambitious multinational projects that aim to increase crop yields by introducing C4 traits into C3 plants. Combined with new technologies, this renewed interest has resulted in the development of more sophisticated approaches toward understanding how the C4 pathway evolved, how it is regulated, and how it might be manipulated. The extent of this resurgence is manifest by the publication in 2011 of more than 650 pages of reviews on different aspects of C4. Here, I provide an overview of our current understanding, the questions that are being addressed, and the issues that lie ahead. PMID:22128120

What does leaf wax δD from a mixed C3/C4 vegetation region tell us?

NASA Astrophysics Data System (ADS)

Wang, Yiming V.; Larsen, Thomas; Leduc, Guillaume; Andersen, Nils; Blanz, Thomas; Schneider, Ralph R.

2013-06-01

Hydrogen isotope values (δD) of sedimentary terrestrial leaf wax such as n-alkanes or n-acids have been used to map and understand past changes in rainfall amount in the tropics because δD of precipitation is commonly assumed as the first order controlling factor of leaf wax δD. Plant functional types and their photosynthetic pathways can also affect leaf wax δD but these biological effects are rarely taken into account in paleo studies relying on this rainfall proxy. To investigate how biological effects may influence δD values we here present a 37,000-year old record of δD and stable carbon isotopes (δ13C) measured on four n-alkanes (n-C27, n-C29, n-C31, n-C33) from a marine sediment core collected off the Zambezi River mouth. Our paleo δ13C records suggest that each individual n-alkanes had different C3/C4 proportional contributions. n-C29 was mostly derived from a C3 dicots (trees, shrubs and forbs) dominant vegetation throughout the entire record. In contrast, the longer chain n-C33 and n-C31 were mostly contributed by C4 grasses during the Glacial period but shifted to a mixture of C4 grasses and C3 dicots during the Holocene. Strong correlations between δD and δ13C values of n-C33 (correlation coefficient R2 = 0.75, n = 58) and n-C31 (R2 = 0.48, n = 58) suggest that their δD values were strongly influenced by changes in the relative contributions of C3/C4 plant types in contrast to n-C29 (R2 = 0.07, n = 58). Within regions with variable C3/C4 input, we conclude that δD values of n-C29 are the most reliable and unbiased indicator for past changes in rainfall, and that δD and δ13C values of n-C31 and n-C33 are sensitive to C3/C4 vegetation changes. Our results demonstrate that a robust interpretation of palaeohydrological data using n-alkane δD requires additional knowledge of regional vegetation changes from which n-alkanes are synthesized, and that the combination of δD and δ13C values of multiple n-alkanes can help to differentiate

Influence of time, temperature and coagulation on the measurement of C3, C3 split products and C4.

PubMed

Sinosich, M J; Teisner, B; Brandslund, I; Fisher, M; Grudzinskas, J G

1982-11-26

Quantitative and qualitative immunoelectrophoretic analyses of circulating C3, C3 split products and C4 were performed in matched EDTA plasma and serum obtained from 5 normal subjects and stored for up to 48 h at room temperature (18 degrees C-22 degrees C) and 4 degrees C. Fluctuations in apparent levels of C3 were greater in serum than plasma stored at room temperature, a fall in levels seen by 24 h being followed by a significant increase. By contrast, levels of C3 did not alter if stored at 4 degrees C. C4 levels in both EDTA plasma and serum remained unchanged for 24 h, a slight decrease being seen at 48 h. Levels of C4 remained constant if samples were stored at 4 degrees C. Crossed immunoelectrophoresis revealed a significant progressive decrease in C3 levels and a simultaneous increase in C3c occurring after 4 h in serum and 8 h in EDTA plasma, stored at room temperature. In studies conducted at 4 degrees C, similar but delayed fluctuations were seen. A progressive and significant increase in C3d levels was seen in both plasma and serum samples stored at room temperature, levels rising to 276% (plasma) and 308% (serum) of levels seen at zero time. At 4 degrees C marginal increases in C3d levels only were observed. These results suggest that in vitro degradation of C3 and C4 are readily facilitated by temperature, time and coagulation, and that conditions of collection and storage of samples must be optimized for the accurate definition of activation of the complement cascade.

Initial events during the evolution of C4 photosynthesis in C3 species of Flaveria.

PubMed

Sage, Tammy L; Busch, Florian A; Johnson, Daniel C; Friesen, Patrick C; Stinson, Corey R; Stata, Matt; Sultmanis, Stefanie; Rahman, Beshar A; Rawsthorne, Stephen; Sage, Rowan F

2013-11-01

The evolution of C4 photosynthesis in many taxa involves the establishment of a two-celled photorespiratory CO2 pump, termed C2 photosynthesis. How C3 species evolved C2 metabolism is critical to understanding the initial phases of C4 plant evolution. To evaluate early events in C4 evolution, we compared leaf anatomy, ultrastructure, and gas-exchange responses of closely related C3 and C2 species of Flaveria, a model genus for C4 evolution. We hypothesized that Flaveria pringlei and Flaveria robusta, two C3 species that are most closely related to the C2 Flaveria species, would show rudimentary characteristics of C2 physiology. Compared with less-related C3 species, bundle sheath (BS) cells of F. pringlei and F. robusta had more mitochondria and chloroplasts, larger mitochondria, and proportionally more of these organelles located along the inner cell periphery. These patterns were similar, although generally less in magnitude, than those observed in the C2 species Flaveria angustifolia and Flaveria sonorensis. In F. pringlei and F. robusta, the CO2 compensation point of photosynthesis was slightly lower than in the less-related C3 species, indicating an increase in photosynthetic efficiency. This could occur because of enhanced refixation of photorespired CO2 by the centripetally positioned organelles in the BS cells. If the phylogenetic positions of F. pringlei and F. robusta reflect ancestral states, these results support a hypothesis that increased numbers of centripetally located organelles initiated a metabolic scavenging of photorespired CO2 within the BS. This could have facilitated the formation of a glycine shuttle between mesophyll and BS cells that characterizes C2 photosynthesis.

Climate controls on C3 vs. C4 productivity in North American grasslands from carbon isotope composition of soil organic matter

USGS Publications Warehouse

von Fischer, J.C.; Tieszen, L.L.; Schimel, D.S.

2008-01-01

We analyzed the ??13 C of soil organic matter (SOM) and fine roots from 55 native grassland sites widely distributed across the US and Canadian Great Plains to examine the relative production of C3 vs. C4 plants (hereafter %C4) at the continental scale. Our climate vs. %C4 results agreed well with North American field studies on %C4, but showed bias with respect to %C4 from a US vegetation database (statsgo) and weak agreement with a physiologically based prediction that depends on crossover temperature. Although monthly average temperatures have been used in many studies to predict %C4, our analysis shows that high temperatures are better predictors of %C4. In particular, we found that July climate (average of daily high temperature and month's total rainfall) predicted %C4 better than other months, seasons or annual averages, suggesting that the outcome of competition between C3 and C4 plants in North American grasslands was particularly sensitive to climate during this narrow window of time. Root ??13 C increased about 1??? between the A and B horizon, suggesting that C 4 roots become relatively more common than C3 roots with depth. These differences in depth distribution likely contribute to the isotopic enrichment with depth in SOM where both C3 and C4 grasses are present. ?? 2008 The Authors Journal compilation ?? 2008 Blackwell Publishing Ltd.

Invasive C4 Perennial Grass Alters Net Ecosystem Exchange in Mixed C3/C4 Savanna Grassland

NASA Astrophysics Data System (ADS)

Basham, T. S.; Litvak, M.

2006-12-01

The invasion of ecosystems by non-native plants that differ from native plants in physiological characteristics and phenology has the potential to alter ecosystem function. In Texas and other regions of the southern central plains of the United States, the introduced C4 perennial grass, Bothriochloa ischaemum, invades C3/C4 mixed grasslands and savannas, resulting in decreased plant community diversity (Gabbard 2003; Harmoney et al 2004). The objective of this study was to quantify how the conversion of these mixed grass communities to C4 dominated, B. ischaemum monocultures impacts carbon cycling and sequestration. Seasonal measurements of Net Ecosystem Exchange (NEE) of CO2, leaf level gas exchange and soil respiration were compared between savanna grassland plots composed of either naturally occurring B. ischaemum monocultures or native mixed grasses (n=16). NEE was measured using a closed system chamber that attached to permanently installed stainless steel bases. Temperature, soil moisture, aerial percent species cover and leaf area index were also monitored in plots to explain variability in measured responses. Results showed that NEE differed seasonally between invaded and native plots due to 1) greater leaf surface area per unit ground area in invaded plots, 2) differences in phenological patterns of plant activity and 3) differences in responses to water limitation between invaded and native plots. Cold season and summer drought NEE were driven primarily by belowground respiration in both plot types, however spring uptake activity commenced two months later in invaded plots. This later start in invaded plots was compensated for by greater uptake throughout the growing season and in particular during the drier summer months. Differences in NEE between plot types were not due to differences in soil respiration nor were they due to greater leaf level photosynthetic capabilities of B. ischaemum relative to the dominant native grasses. NEE, soil respiration and

Major alterations in transcript profiles between C3-C4 and C4 photosynthesis of an amphibious species Eleocharis baldwinii.

PubMed

Chen, Taiyu; Zhu, Xin-Guang; Lin, Yongjun

2014-09-01

Engineering C4 photosynthetic metabolism into C3 crops is regarded as a major strategy to increase crop productivity, and clarification of the evolutionary processes of C4 photosynthesis can help the better use of this strategy. Here, Eleocharis baldwinii, a species in which C4 photosynthesis can be induced from a C3-C4 state under either environmental or ABA treatments, was used to identify the major transcriptional modifications during the process from C3-C4 to C4. The transcriptomic comparison suggested that in addition to the major differences in C4 core pathway, the pathways of glycolysis, citrate acid metabolism and protein synthesis were dramatically modified during the inducement of C4 photosynthetic states. Transcripts of many transporters, including not only metabolite transporters but also ion transporters, were dramatically increased in C4 photosynthetic state. Many candidate regulatory genes with unidentified functions were differentially expressed in C3-C4 and C4 photosynthetic states. Finally, it was indicated that ABA, auxin signaling and DNA methylation play critical roles in the regulation of C4 photosynthesis. In summary, by studying the different photosynthetic states of the same species, this work provides the major transcriptional differences between C3-C4 and C4 photosynthesis, and many of the transcriptional differences are potentially related to C4 development and therefore are the potential targets for reverse genetics studies.

Little Cross-Feeding of the Mycorrhizal Networks Shared Between C3-Panicum bisulcatum and C4-Panicum maximum Under Different Temperature Regimes

PubMed Central

Řezáčová, Veronika; Zemková, Lenka; Beskid, Olena; Püschel, David; Konvalinková, Tereza; Hujslová, Martina; Slavíková, Renata; Jansa, Jan

2018-01-01

Common mycorrhizal networks (CMNs) formed by arbuscular mycorrhizal fungi (AMF) interconnect plants of the same and/or different species, redistributing nutrients and draining carbon (C) from the different plant partners at different rates. Here, we conducted a plant co-existence (intercropping) experiment testing the role of AMF in resource sharing and exploitation by simplified plant communities composed of two congeneric grass species (Panicum spp.) with different photosynthetic metabolism types (C3 or C4). The grasses had spatially separated rooting zones, conjoined through a root-free (but AMF-accessible) zone added with 15N-labeled plant (clover) residues. The plants were grown under two different temperature regimes: high temperature (36/32°C day/night) or ambient temperature (25/21°C day/night) applied over 49 days after an initial period of 26 days at ambient temperature. We made use of the distinct C-isotopic composition of the two plant species sharing the same CMN (composed of a synthetic AMF community of five fungal genera) to estimate if the CMN was or was not fed preferentially under the specific environmental conditions by one or the other plant species. Using the C-isotopic composition of AMF-specific fatty acid (C16:1ω5) in roots and in the potting substrate harboring the extraradical AMF hyphae, we found that the C3-Panicum continued feeding the CMN at both temperatures with a significant and invariable share of C resources. This was surprising because the growth of the C3 plants was more susceptible to high temperature than that of the C4 plants and the C3-Panicum alone suppressed abundance of the AMF (particularly Funneliformis sp.) in its roots due to the elevated temperature. Moreover, elevated temperature induced a shift in competition for nitrogen between the two plant species in favor of the C4-Panicum, as demonstrated by significantly lower 15N yields of the C3-Panicum but higher 15N yields of the C4-Panicum at elevated as compared to

Comparison of carbon balance in Mediterranean pilot constructed wetlands vegetated with different C4 plant species.

PubMed

Barbera, Antonio C; Borin, Maurizio; Cirelli, Giuseppe L; Toscano, Attilio; Maucieri, Carmelo

2015-02-01

This study investigates carbon dioxide (CO2) and methane (CH4) emissions and carbon (C) budgets in a horizontal subsurface flow pilot-plant constructed wetland (CW) with beds vegetated with Cyperus papyrus L., Chrysopogon zizanioides (L.) Roberty, and Mischantus × giganteus Greef et Deu in the Mediterranean basin (Sicily) during the 1st year of plant growing season. At the end of the vegetative season, M. giganteus showed the higher biomass accumulation (7.4 kg m(-2)) followed by C. zizanioides (5.3 kg m(-2)) and C. papyrus (1.8 kg m(-2)). Significantly higher emissions of CO2 were detected in the summer, while CH4 emissions were maximum during spring. Cumulative CO2 emissions by C. papyrus and C. zizanioides during the monitoring period showed similar trends with final values of about 775 and 1,074 g m(-2), respectively, whereas M. giganteus emitted 3,395 g m(-2). Cumulative CH4 bed emission showed different trends for the three C4 plant species in which total gas release during the study period was for C. papyrus 12.0 g m(-2) and ten times higher for M. giganteus, while C. zizanioides bed showed the greatest CH4 cumulative emission with 240.3 g m(-2). The wastewater organic carbon abatement determined different C flux in the atmosphere. Gas fluxes were influenced both by plant species and monitored months with an average C-emitted-to-C-removed ratio for C. zizanioides, C. papyrus, and M. giganteus of 0.3, 0.5, and 0.9, respectively. The growing season C balances were positive for all vegetated beds with the highest C sequestered in the bed with M. giganteus (4.26 kg m(-2)) followed by C. zizanioides (3.78 kg m(-2)) and C. papyrus (1.89 kg m(-2)). To our knowledge, this is the first paper that presents preliminary results on CO2 and CH4 emissions from CWs vegetated with C4 plant species in Mediterranean basin during vegetative growth.

Stable C &N isotopes in 2100 Year-B.P. human bone collagen indicate rare dietary dominance of C4 plants in NE-Italy.

PubMed

Laffranchi, Zita; Huertas, Antonio Delgado; Jiménez Brobeil, Sylvia A; Torres, Arsenio Granados; Riquelme Cantal, Jose A

2016-12-09

C 4 plants (e.g. maize, millet), part of our current diet, are only endemic of reduced areas in South-Europe due to their need of warm climates. Since the first vestiges of agriculture in Europe remains of C 4 plants were recorded but their overall proportion in the human diet remains unknown. Therefore, isotopic (δ 13 C and δ 15 N) composition of bone collagen from the skeletal remains (human and animals) of a Celtic population, Cenomani Gauls, from Verona (3 rd to 1 st century BC) in the NE Italy provide a new perspective on this matter. The δ 13 C collagen values of 90 human skeletal individuals range between -20.2‰ and -9.7‰ (V-PDB) with a mean value of -15.3‰. As present day C 4 plants have δ 13 C values around -11‰, which is equivalent to -9.5‰ for samples of preindustrial age, the less negative δ 13 C values in these individuals indicate a diet dominated by C 4 plants. This palaeodietary study indicates that some European populations predominantly consumed cultivated C 4 plants 2100 year B.P. This is supported by the paleobotanical records and ancient Roman sources (e.g. Pliny the Elder), which indicate that millet was a staple food in South-Europe.

Stable C & N isotopes in 2100 Year-B.P. human bone collagen indicate rare dietary dominance of C4 plants in NE-Italy

NASA Astrophysics Data System (ADS)

Laffranchi, Zita; Huertas, Antonio Delgado; Jiménez Brobeil, Sylvia A.; Torres, Arsenio Granados; Riquelme Cantal, Jose A.

2016-12-01

C4 plants (e.g. maize, millet), part of our current diet, are only endemic of reduced areas in South-Europe due to their need of warm climates. Since the first vestiges of agriculture in Europe remains of C4 plants were recorded but their overall proportion in the human diet remains unknown. Therefore, isotopic (δ13C and δ15N) composition of bone collagen from the skeletal remains (human and animals) of a Celtic population, Cenomani Gauls, from Verona (3rd to 1st century BC) in the NE Italy provide a new perspective on this matter. The δ13C collagen values of 90 human skeletal individuals range between -20.2‰ and -9.7‰ (V-PDB) with a mean value of -15.3‰. As present day C4 plants have δ13C values around -11‰, which is equivalent to -9.5‰ for samples of preindustrial age, the less negative δ13C values in these individuals indicate a diet dominated by C4 plants. This palaeodietary study indicates that some European populations predominantly consumed cultivated C4 plants 2100 year B.P. This is supported by the paleobotanical records and ancient Roman sources (e.g. Pliny the Elder), which indicate that millet was a staple food in South-Europe.

Stable C & N isotopes in 2100 Year-B.P. human bone collagen indicate rare dietary dominance of C4 plants in NE-Italy

PubMed Central

Laffranchi, Zita; Huertas, Antonio Delgado; Jiménez Brobeil, Sylvia A.; Torres, Arsenio Granados; Riquelme Cantal, Jose A.

2016-01-01

C4 plants (e.g. maize, millet), part of our current diet, are only endemic of reduced areas in South-Europe due to their need of warm climates. Since the first vestiges of agriculture in Europe remains of C4 plants were recorded but their overall proportion in the human diet remains unknown. Therefore, isotopic (δ13C and δ15N) composition of bone collagen from the skeletal remains (human and animals) of a Celtic population, Cenomani Gauls, from Verona (3rd to 1st century BC) in the NE Italy provide a new perspective on this matter. The δ13C collagen values of 90 human skeletal individuals range between −20.2‰ and −9.7‰ (V-PDB) with a mean value of −15.3‰. As present day C4 plants have δ13C values around −11‰, which is equivalent to −9.5‰ for samples of preindustrial age, the less negative δ13C values in these individuals indicate a diet dominated by C4 plants. This palaeodietary study indicates that some European populations predominantly consumed cultivated C4 plants 2100 year B.P. This is supported by the paleobotanical records and ancient Roman sources (e.g. Pliny the Elder), which indicate that millet was a staple food in South-Europe. PMID:27934943

Interfacial coupling induced direct Z scheme water splitting in metal-free photocatalyst: C3N/g-C3N4 heterojunctions.

PubMed

Wang, Jiajun; Li, Xiaoting; You, Ya; Xintong, Yang; Wang, Ying; Li, Qunxiang

2018-06-21

Mimicking the natural photosynthesis in green plants, artificial Z-scheme photocatalysis enables more efficient utilization of solar energy for photocatalytic water splitting. Most currently designed g-C3N4-based Z-scheme heterojunctions are usually based on metal-containing semiconductor photocatalysts, thus exploiting metal-free photocatalysts for Z-scheme water splitting is of huge interest. Herein, we propose two metal-free C3N/g-C3N4 heterojunctions with the C3N monolayer covering g-C3N4 sheet (monolayer or bilayer) and systematically explore their electronic structures, charge distributions and photocatalytic properties by performing extensive hybrid density functional calculations. We clearly reveal that the relative strong built-in electric fields around their respective interface regions, caused by the charge transfer from C3N monolayer to g-C3N4 monolayer or bilayer, result in the bands bending, renders the transfer of photogenerated carriers in these two heterojunctions following the Z-scheme instead of the type-II pathway. Moreover, the photogenerated electrons and holes in these two C3N/g-C3N4 heterojunctions not only can be efficiently separated, but also have strong redox abilities for water oxidation and reduction. Compared with the isolated g-C3N4 sheets, the light absorption in visible to near-infrared region are significantly enhanced in these proposed heterojunctions. These theoretical findings suggest that these proposed metal-free C3N/g-C3N4 heterojunctions are promising direct Z-scheme photocatalysts for solar water splitting. © 2018 IOP Publishing Ltd.

Climate controls on savanna C3 and C4 expansion in Southern Africa during the last 36 kyr BP

NASA Astrophysics Data System (ADS)

Wang, Y. V.; Larsen, T.; Andersen, N.; Blanz, T.; Schneider, R. R.

2010-12-01

Savannahs contain a mixture of C3 and C4 vegetation, accounting for more than a quarter of global primary production and are the second most important biome on the continents. However, our understanding on how savannahs will respond to rising CO2 concentration and temperatures or the IPCC estimated decrease in rainfall is not yet clear in spite of potential far reaching socio-economic consequences. In this study, we used the δD and δ13C of sedimentary long-chain n-alkanes (n-C27,29,31,33 ) in concert with reconstructions for sea surface temperatures and fluvial discharge from a marine sediment core (GIK16160-3, 18°14.47’S, 37°52.27’W, 1334m water depth), collected near the Zambezi river mouth to examine savannah responses under different hydrological and climate conditions in Southern Africa during the last 36 kyr BP. Our data show large variability in both δD and δ13C records of the four n-alkanes, with isotopic differences between individual n-alkanes being far more pronounced during the Glacial than during the Deglacial and Holocene. These large differences may be explained by proportionally higher contributions of C4 grasses over C3 trees to the n-C33,31, which seems to be opposite for n-C29. A strong anticorrelation between δD and δ13C from 36 to 16 kyr BP for n-C31 (R2=0.55) and n-C33 (R2=0.70) suggests that δD of these n-alkanes is strongly influenced by changes in vegetation types as well as physiological effects, rather than being directly related to evaporation/ precipitation balance. In contrast, no apparent relationship (R2=0.32) exists between δD and δ13C of n-C29, suggesting that n-C29 is the most promising hydrological proxy due to less variable vegetation type contributions to n-C29 throughout the core. The C4 plant contribution, which was estimated by taking into account the four n-alkanes δ13C signals and their abundance, implies dominance of C4 grass between 36 and 20 kyr BP, and more evenly distributed C3 and C4 vegetation from

Effects of climate and water balance across grasslands of varying C3 and C4 grass cover

USGS Publications Warehouse

Witwicki, Dana L.; Munson, Seth M.; Thoma, David P.

2016-01-01

Climate change in grassland ecosystems may lead to divergent shifts in the abundance and distribution of C3 and C4 grasses. Many studies relate mean climate conditions over relatively long time periods to plant cover, but there is still much uncertainty about how the balance of C3and C4 species will be affected by climate at a finer temporal scale than season (individual events to months). We monitored cover at five grassland sites with co-dominant C3 and C4 grass species or only dominant C3 grass species for 6 yr in national parks across the Colorado Plateau region to assess the influence of specific months of climate and water balance on changes in grass cover. C4 grass cover increased and decreased to a larger degree than C3 grass cover with extremely dry and wet consecutive years, but this response varied by ecological site. Climate and water balance explained 10–49% of the inter-annual variability of cover of C3 and C4 grasses at all sites. High precipitation in the spring and in previous year monsoon storms influenced changes in cover of C4 grasses, with measures of water balance in the same months explaining additional variability. C3 grasses in grasslands where they were dominant were influenced primarily by longer periods of climate, while C3 grasses in grasslands where they were co-dominant with C4 grasses were influenced little by climate anomalies at either short or long periods of time. Our results suggest that future changes in spring and summer climate and water balance are likely to affect cover of both C3 and C4 grasses, but cover of C4 grasses may be affected more strongly, and the degree of change will depend on soils and topography where they are growing and the timing of the growing season.

Investigations of 3C-SiC inclusions in 4H-SiC epilayers on 4H-SiC single crystal substrates

DOE Office of Scientific and Technical Information (OSTI.GOV)

Si, W.; Dudley, M.; Kong, H.S.

1997-03-01

Synchrotron white beam x-ray topography (SWBXT) and Nomarski optical microscopy (NOM) have been used to characterize 4H-SiC epilayers and to study the character of triangular inclusions therein. 4H-SiC substrates misoriented by a range of angles from (0001), as well as (1 1{bar 0}0) and (11 2{bar 0}) oriented substrates were used. No evidence was found for the nucleation of 3C-SiC inclusions at superscrew dislocations (along the [0001] axis) in the 4H-SiC substrates. Increasing the off-axis angle of the substrates from 3.5 to 6.5{degree} was found to greatly suppress the formation of the triangular inclusions. In the case of substrates misorientedmore » by 8.0{degree} from (0001) toward [112{bar 0}], the triangular inclusions were virtually eliminated. The crystalline quality of 4H-SiC epilayers grown on the substrates misoriented by 8.0{degree} from (0001) was very good. For the (11{bar 0}0) and (112{bar 0}) samples, there is no indication of 3C-SiC inclusions in the epilayers. Possible formation mechanisms and the morphology of 3C-SiC inclusions are discussed. 17 refs., 13 figs.« less

CFD Growth of 3C-SiC on 4H/6H Mesas

NASA Technical Reports Server (NTRS)

Neudeck, Philip G.; Trunek, Andrew J.; Spry, David J.; Powell, J. Anthony; Du, Hui; Skowronski, Marek; Huang, XianRong; Dudley, Michael

2006-01-01

This article describes growth and characterization of the highest quality reproducible 3C-SiC heteroepitaxial films ever reported. By properly nucleating 3C-SiC growth on top of perfectly on-axis (0001) 4H-SiC mesa surfaces completely free of atomic scale steps and extended defects, growth of 3C-SiC mesa heterofilms completely free of extended crystal defects can be achieved. In contrast, nucleation and growth of 3C-SiC mesa heterofilms on top of 4H-SiC mesas with atomic-scale steps always results in numerous observable dislocations threading through the 3C-SiC epilayer. High-resolution X-ray diffraction and transmission electron microscopy measurements indicate non-trivial in-plane lattice mismatch between the 3C and 4H layers. This mismatch is somewhat relieved in the step-free mesa case via misfit dislocations confined to the 3C/4H interfacial region without dislocations threading into the overlying 3C-SiC layer. These results indicate that the presence or absence of steps at the 3C/4H heteroepitaxial interface critically impacts the quality, defect structure, and relaxation mechanisms of single-crystal heteroepitaxial 3C-SiC films.

Copper(I)-catalyzed substitution reactions of propargylic amines: importance of C(sp)-C(sp3) bond cleavage in generation of iminium intermediates.

PubMed

Sugiishi, Tsuyuka; Kimura, Akifumi; Nakamura, Hiroyuki

2010-04-21

Substitution reactions of propargylic amines proceed in the presence of copper(I) catalysts. Mechanistic studies showed that C(sp)-C(sp(3)) bond cleavage assisted by nitrogen lone-pair electrons is essential for the reaction, and the resulting iminium intermediates undergo amine exchange, aldehyde exchange, and alkyne addition reactions. Because iminium intermediates are key to aldehyde-alkyne-amine (A(3)) coupling reactions, this transformation is effective not only for reconstruction of propargylic amines but also for chiral induction of racemic compounds in the presence of chiral catalysts.

Carbon storage potential increases with increasing ratio of C4 to C3 grass cover and soil productivity in restored tallgrass prairies.

PubMed

Spiesman, Brian J; Kummel, Herika; Jackson, Randall D

2018-02-01

Long-term soil carbon (C) storage is essential for reducing CO 2 in the atmosphere. Converting unproductive and environmentally sensitive agricultural lands to grasslands for bioenergy production may enhance C storage. However, a better understanding of the interacting effects of grass functional composition (i.e., relative abundance of C 4 and C 3 grass cover) and soil productivity on C storage will help guide sustainable grassland management. Our objective was to examine the relationship between grass functional composition and potential C storage and how it varies with potential soil productivity. We estimated C inputs from above- and belowground net primary productivity (ANPP and BNPP), and heterotrophic respiration (R H ) to calculate net ecosystem production (NEP), a measure of potential soil C storage, in grassland plots of relatively high- and low-productivity soils spanning a gradient in the ratio of C 4 to C 3 grass cover (C 4 :C 3 ). NEP increased with increasing C 4 :C 3 , but only in potentially productive soils. The positive relationship likely stemmed from increased ANPP, rather than BNPP, which was possibly related to efficient resource-use and physiological/anatomical advantages of C 4 plants. R H was negatively correlated with C 4 :C 3 , possibly because of changes in microclimate or plant-microbe interactions. It is possible that in potentially productive soils, C storage can be enhanced by favoring C 4 over C 3 grasses through increased ANPP and BNPP and reduced R H . Results also suggest that potential C storage gains from C 4 productivity would not be undermined by a corresponding increase in R H .

Primary Glomerulonephritis with Unique C4d Deposition and Concurrent Non-infectious Intermediate Uveitis: a Case Report and Literature Review

PubMed Central

2018-01-01

C4 glomerulopathy is a recently introduced entity that presents with bright C4d staining and minimal or absent immunoglobulin and C3 staining. We report a case of a 62-year-old man with C4 glomerulonephritis (GN) and uveitis. He presented to the nephrology department with proteinuria and hematuria. The patient also had intermediate uveitis along with proteinuria and hematuria. A kidney biopsy that was performed in light of continuing proteinuria and hematuria showed a focal proliferative, focal sclerotic glomerulopathy pattern on light microscopy, absent staining for immunoglobulin or C3 by immunofluorescence microscopy, with bright staining for C4d on immunohistochemistry, and electron-dense deposits on electron microscopy. Consequently, C4 GN was suggested as the pathologic diagnosis. Although laser microdissection and mass spectrometry for glomerular deposit and pathologic evaluation of the retinal tissue were not performed, this is the first report of C4 GN in Korea and the first case of coexisting C4 GN and uveitis in the English literature. PMID:29713256

Biocatalytic route to C-3'-azido/-hydroxy-C-4'-spiro-oxetanoribonucleosides.

PubMed

Kumar, Manish; Sharma, Vivek K; Kumar, Rajesh; Prasad, Ashok K

2015-11-19

The lipase, Novozyme(®)-435, exclusively deacetylates the 5-O-acetyl over 4-C-acetyloxymethyl group of almost identical reactivity in 5-O-acetyl-4-C-acetyloxymethyl-3-azido-3-deoxy-1,2-O-isopropylidene-α-D-ribofuranose that led to the development of first and efficient synthesis of 3'-azido-/3'-amino-C-4'-spiro-oxetanoribonucleosides T, U, C and A in 20-24% overall yields. The X-ray study on the compound obtained by tosylation of lipase-mediated monodeacetylated product unambiguously confirmed the point of diastereoselective monodeacetylation on diacetoxy-azido-ribofuranose derivative. The capability of Novozyme(®)-435 for selective deacylation of 5-O-acetyl group in 5-O-acetyl-4-C-acetyloxymethyl-3-O-benzyl-1,2-O-isopropylidene-α-D-ribofuranose recently discovered by us has been successfully used for the synthesis of C-4'-spiro-oxetanoribonucleosides A and C in good yields. These results clearly indicate that the broader substrate specificity and highly selective capability of Novozyme(®)-435 for carrying out acetylation/deacetylation reactions can be utilized for the development of environment friendly selective methodologies in organic synthesis. Copyright © 2015 Elsevier Ltd. All rights reserved.

Carbon isotope compositions of terrestrial C3 plants as indicators of (paleo)ecology and (paleo)climate.

PubMed

Kohn, Matthew J

2010-11-16

A broad compilation of modern carbon isotope compositions in all C3 plant types shows a monotonic increase in δ(13)C with decreasing mean annual precipitation (MAP) that differs from previous models. Corrections for temperature, altitude, or latitude are smaller than previously estimated. As corrected for altitude, latitude, and the δ(13)C of atmospheric CO(2), these data permit refined interpretation of MAP, paleodiet, and paleoecology of ecosystems dominated by C3 plants, either prior to 7-8 million years ago (Ma), or more recently at mid- to high latitudes. Twenty-nine published paleontological studies suggest preservational or scientific bias toward dry ecosystems, although wet ecosystems are also represented. Unambiguous isotopic evidence for C4 plants is lacking prior to 7-8 Ma, and hominid ecosystems at 4.4 Ma show no isotopic evidence for dense forests. Consideration of global plant biomass indicates that average δ(13)C of C3 plants is commonly overestimated by approximately 2‰.

Prospects for improving CO2 fixation in C3-crops through understanding C4-Rubisco biogenesis and catalytic diversity.

PubMed

Sharwood, Robert E; Ghannoum, Oula; Whitney, Spencer M

2016-06-01

By operating a CO2 concentrating mechanism, C4-photosynthesis offers highly successful solutions to remedy the inefficiency of the CO2-fixing enzyme Rubisco. C4-plant Rubisco has characteristically evolved faster carboxylation rates with low CO2 affinity. Owing to high CO2 concentrations in bundle sheath chloroplasts, faster Rubisco enhances resource use efficiency in C4 plants by reducing the energy and carbon costs associated with photorespiration and lowering the nitrogen investment in Rubisco. Here, we show that C4-Rubisco from some NADP-ME species, such as maize, are also of potential benefit to C3-photosynthesis under current and future atmospheric CO2 pressures. Realizing this bioengineering endeavour necessitates improved understanding of the biogenesis requirements and catalytic variability of C4-Rubisco, as well as the development of transformation capabilities to engineer Rubisco in a wider variety of food and fibre crops. Copyright © 2016 Elsevier Ltd. All rights reserved.

Isotropic C6, C8 and C10 interaction coefficients for CH 4, C 2H 6, C 3H 8, n-C 4H 10 and cyclo- C3H 6

NASA Astrophysics Data System (ADS)

Thomas, Gerald F.; Mulder, Fred; Meath, William J.

1980-12-01

The non-empirical generalized Kirkwood, Unsöld, and the single-Δ Unsöld methods (with double-zeta quality SCF wave-functions) are used to calculate isotropic dispersion (and induction) energy coefficients C2n, with n ⩽ 5, for interactions involving ground state CH 4, C 2H 6, C 3H 8, n-C 4H 10 and cyclo-C 3H 6. Results are also given for the related multipole polarizabilities α l, multipole sums S1/(0) and S1(-1) which are evaluated using sum rules, and the permanent multipole moments. for l = 1 (dipole) to l = 3 (octupole). Estimates of the reliability of the non-empirical methods, for the type of molecules considered, are obtained by a comparison with accurate literature values of α 1S1(-1) and C6. This, and the asymptotic properties of the multipolar expansion of the dispersion energy, the use to discuss recommended representation for the isotropic long range interaction energies through R-10 where R is the intermolecular separation.

Timing and Nature of Appearance of C4 Plants in the Indian Subcontinent: Clue from Isotopic Ratios of Biomarker

NASA Astrophysics Data System (ADS)

Sanyal, P.

2014-12-01

Appearance and expansion of C4 plants during the late Miocene was first documented from Siwalik Group of sediments using carbon isotope ratio of soil carbonates. It was proposed that C4 plants appeared drastically in response to change in hydrological cycle. Subsequently, various workers documented evolution and expansion of C4 plants from different Siwalik sections of India, Nepal and Pakistan. Although evolution and expansion of C4 plants was documented in all these studies but nature and timing of expansion of C4 plants showed variations from section to sections. Even from same section, the results varied with change in proxies. The hydrological changes also differ regionally. One of the reasons of differences in results was lack of modern data set from the Indian subcontinent. For example, in many cases δ18O values of New Delhi rainwater has been considered as reference data set and applied to site of investigation with some corrections. The average δ13C values of C3 and C4 plants was considered as -27‰ and -12.5‰ respectively, but modern C3 plants in the Gangetic plain is almost 2.5‰ lower than the value used in those studies. Additional problem raised as preservation of pristine isotopic character of soil organic matter is also apprehensive. To resolve all these issues, the δ13C value of long chain alkane have been used to reconstruct C3-C4 plants after isotopic characterization of modern plants and their biomarker from the Gangetic plain which is equivalent to the past Siwalik flood plain. Additionally, δD values of long chain alkane were also measured. Variations of d13C values of long chain alkane with time show presence of C4 plants in Siwalik from 11 Ma ago and since then gradual increase in C4 plants till 6 Ma followed by a sharp increase. The statistically significant correlation between δD and δ13C values of long chain alkane show that positive impact of hydrological change on abundance of C3-C4 plants.

Heat inactivation of leaf phosphoenolpyruvate carboxylase: Protection by aspartate and malate in C4 plants.

PubMed

Rathnam, C K

1978-01-01

The activity of phosphoenolpyruvate (PEP) carboxylase EC 4.1.1.31 in leaf extracts of Eleusine indica L. Gaertn., a C4 plant, exhibited a temperature optimum of 35-37° C with a complete loss of activity at 50° C. However, the enzyme was protected effectively from heat inactivation up to 55° C by L-aspartate. Activation energies (Ea) for the enzyme in the presence of aspartate were 2.5 times lower than that of the control enzyme. Arrhenius plots of PEP carboxylase activity (±aspartate) showed a break in the slope around 17-20° C with a 3-fold increase in the Ea below the break. The discontinuity in the slopes was abolished by treating the enzyme extracts with Triton X-100, suggesting that PEP carboxylase in C4 plants is associated with lipid and may be a membrane bound enzyme. Depending upon the species, the major C4 acid formed during photosynthesis (malate or aspartate) was found to be more protective than the minor C4 acid against the heat inactivation of their PEP carboxylase. Oxaloacetate, the reaction product, was less effective compared to malate or aspartate. Several allosteric inhibitors of PEP carboxylase were found to be moderately to highly effective in protecting the C4 enzyme while its activators showed no significant effect. PEP carboxylase from C3 species was not protected from thermal inactivation by the C4 acids. The physiological significance of these results is discussed in relation to the high temperature tolerance of C4 plants.

Tracking the evolutionary rise of C4 metabolism.

PubMed

Sage, Rowan F

2016-05-01

Upregulation of the C4 metabolic cycle is a major step in the evolution of C4 photosynthesis. Why this happened remains unclear, in part because of difficulties measuring the C4 cycle in situ in C3-C4 intermediate species. Now, Alonso-Cantabrana and von Caemmerer (2016) have described a new approach for quantifying C4 cycle activity, thereby providing the means to analyze its upregulation in an evolutionary context. © The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology.

A critical review on the improvement of photosynthetic carbon assimilation in C3 plants using genetic engineering.

PubMed

Ruan, Cheng-Jiang; Shao, Hong-Bo; Teixeira da Silva, Jaime A

2012-03-01

Global warming is one of the most serious challenges facing us today. It may be linked to the increase in atmospheric CO2 and other greenhouse gases (GHGs), leading to a rise in sea level, notable shifts in ecosystems, and in the frequency and intensity of wild fires. There is a strong interest in stabilizing the atmospheric concentration of CO2 and other GHGs by decreasing carbon emission and/or increasing carbon sequestration. Biotic sequestration is an important and effective strategy to mitigate the effects of rising atmospheric CO2 concentrations by increasing carbon sequestration and storage capacity of ecosystems using plant photosynthesis and by decreasing carbon emission using biofuel rather than fossil fuel. Improvement of photosynthetic carbon assimilation, using transgenic engineering, potentially provides a set of available and effective tools for enhancing plant carbon sequestration. In this review, firstly different biological methods of CO2 assimilation in C3, C4 and CAM plants are introduced and three types of C4 pathways which have high photosynthetic performance and have evolved as CO2 pumps are briefly summarized. Then (i) the improvement of photosynthetic carbon assimilation of C3 plants by transgenic engineering using non-C4 genes, and (ii) the overexpression of individual or multiple C4 cycle photosynthetic genes (PEPC, PPDK, PCK, NADP-ME and NADP-MDH) in transgenic C3 plants (e.g. tobacco, potato, rice and Arabidopsis) are highlighted. Some transgenic C3 plants (e.g. tobacco, rice and Arabidopsis) overexpressing the FBP/SBPase, ictB and cytochrome c6 genes showed positive effects on photosynthetic efficiency and growth characteristics. However, over the last 28 years, efforts to overexpress individual, double or multiple C4 enzymes in C3 plants like tobacco, potato, rice, and Arabidopsis have produced mixed results that do not confirm or eliminate the possibility of improving photosynthesis of C3 plants by this approach. Finally, a prospect

Carbon isotope compositions of terrestrial C3 plants as indicators of (paleo)ecology and (paleo)climate

PubMed Central

Kohn, Matthew J.

2010-01-01

A broad compilation of modern carbon isotope compositions in all C3 plant types shows a monotonic increase in δ13C with decreasing mean annual precipitation (MAP) that differs from previous models. Corrections for temperature, altitude, or latitude are smaller than previously estimated. As corrected for altitude, latitude, and the δ13C of atmospheric CO2, these data permit refined interpretation of MAP, paleodiet, and paleoecology of ecosystems dominated by C3 plants, either prior to 7–8 million years ago (Ma), or more recently at mid- to high latitudes. Twenty-nine published paleontological studies suggest preservational or scientific bias toward dry ecosystems, although wet ecosystems are also represented. Unambiguous isotopic evidence for C4 plants is lacking prior to 7–8 Ma, and hominid ecosystems at 4.4 Ma show no isotopic evidence for dense forests. Consideration of global plant biomass indicates that average δ13C of C3 plants is commonly overestimated by approximately 2‰. PMID:21041671

Influence of water stress on the carbon isotopic composition of modern plants: Implications for C4 plant appearance in Indian subcontinent

NASA Astrophysics Data System (ADS)

Basu, S.; Ghosh, S.; Sanyal, P.

2016-12-01

The carbon isotopic composition (d13C) of modern terrestrial plants (C3 and C4) provides the baseline to understand past vegetation composition, paleodietary changes and animal migration etc. Accuracy of past environment reconstruction is dependent on the end-member d13C values of plants which found to vary in regional scale. For instance, the d13C values of Indian C3 plants (d13CC3) are 1 to 2‰ more negative compared to global mean. As observed, most of the previous database is devoid of samples from tropical monsoon realm (like India) and the difference between global and regional mean may introduce errors in vegetation reconstruction. To constrain end-member d13CC3 value, published and newly generated results from wide range of mean annual precipitation (MAP: 1-11,700 mm) are compiled which is ca. 1.5 higher in sample size (n=2440) compared to previous database. Using logarithmic function, new relationship between d13C value and MAP (d13CC3 (‰) =20.1585(0.3061)-1.1276(0.0489)ln(MAP+700)) is proposed. The modeled mean d13CC3 value (-28.9‰) is close to average d13CC3 value for Indian plants (-29.1‰) and suggests the importance of vegetation from low-latitudinal tropical region in global compilation. It was observed that C3 plants, on a global scale, are less sensitive to wet climate relative to dry condition. This inference is in agreement with the paleoclimatic data from Indian subcontinent for the late Quaternary period. Despite well established correlation between d13CC3 value and MAP, previous investigation from Indian subcontinent used fixed end-member values to reconstruct past vegetation and total change in the d13C value of proxies was attributed to changes in relative abundance of C3 and C4 plants. Using region-specific mean d13C value of plants, after correcting for changing MAP and error propagation, existence of C4 before ca. 11 Ma plant is observed; earlier to previous reported timing.

Induction of kranz anatomy and C4-like biochemical characteristics in a submerged amphibious plant by abscisic acid

PubMed Central

Ueno, O

1998-01-01

The amphibious leafless sedge Eleocharis vivipara develops C4-like traits as well as Kranz anatomy under terrestrial conditions, but it develops C3-like traits without Kranz anatomy under submerged conditions. When submerged plants are exposed to aerial conditions, they rapidly produce new photosynthetic tissues with C4-like traits. In this study, experiments were performed to determine whether abscisic acid (ABA), a plant stress hormone, could induce the formation of photosynthetic tissues with Kranz anatomy and C4-like biochemical traits under water in the submerged form. When the submerged plants were grown in water containing 5 &mgr;M ABA, they developed new photosynthetic tissues with Kranz anatomy, forming well-developed Kranz (bundle sheath) cells that contained many organelles. The ABA-induced tissues accumulated large amounts of phosphoenolpyruvate carboxylase, pyruvate orthophosphate dikinase, and NAD-malic enzyme at the appropriate cellular sites. The tissues had 3.4 to 3.8 times more C4 enzyme activity than did tissues of the untreated submerged plants. Carbon-14 pulse and carbon-12 chase experiments revealed that the ABA-induced tissues fixed higher amounts of carbon-14 into C4 compounds and lower amounts of carbon-14 into C3 compounds as initial products than did the submerged plants and that they exhibited a C4-like pattern of carbon fixation under aqueous conditions of low carbon, indicating enhanced C4 capacity in the tissues. This report provides an example of the hormonal control of the differentiation of the structural and functional traits required for the C4 pathway. PMID:9548983

Effect of 3C-SiC intermediate layer in GaN—based light emitting diodes grown on Si(111) substrate

NASA Astrophysics Data System (ADS)

Zhu, Youhua; Wang, Meiyu; Li, Yi; Tan, Shuxin; Deng, Honghai; Guo, Xinglong; Yin, Haihong; Egawa, Takashi

2017-03-01

GaN-based light emitting diodes (LEDs) have been grown by metalorganic chemical vapor deposition on Si(111) substrate with and without 3C-SiC intermediate layer (IL). Structural property has been characterized by means of atomic force microscope, X-ray diffraction, and transmission electron microscope measurements. It has been revealed that a significant improvement in crystalline quality of GaN and superlattice epitaxial layers can be achieved by using 3C-SiC as IL. Regarding of electrical and optical characteristics, it is clearly observed that the LEDs with its IL have a smaller leakage current and higher light output power comparing with the LEDs without IL. The better performance of LEDs using 3C-SiC IL can be contributed to both of the improvements in epitaxial layers quality and light extraction efficiency. As a consequence, in terms of optical property, a double enhancement of the light output power and external quantum efficiency has been realized.

Differential freezing resistance and photoprotection in C3 and C4 eudicots and grasses

PubMed Central

Liu, Mei-Zhen; Osborne, Colin P.

2013-01-01

Globally, C4 plants dominate hot, open environments, but this general pattern is underpinned by important differences in the biogeography of C4 lineages. In particular, the species richness of C4 Poaceae (grasses) increases strongly with increasing temperature, whereas that of the major C4 eudicot group Chenopodiaceae correlates positively with aridity. Freezing tolerance is a crucial determinant of biogeographical relationships with temperature and is mediated by photodamage and cellular disruption by desiccation, but little is known about differences between C4 families. This study hypothesized that there is a greater risk of freezing damage via these mechanisms in C4 Poaceae than Chenopodiaceae, that freezing protection differs between the taxonomic groups, and that freezing tolerance of species is linked to arid habitat preference. Chlorophyll fluorescence, water relations, and freezing injury were compared in four C3 and six C4 species of Poaceae and Chenopodiaceae from the same Mongolian flora. Contrary to expectations, freezing-induced leaf mortality and photodamage were lower in Poaceae than Chenopodiaceae species, and unrelated to photosynthetic pathway. The freezing resistance of Poaceae species resulted from constitutive protection and cold acclimation and an ability to protect the photosynthetic apparatus from photodamage. Freezing protection was associated with low osmotic potential and low tissue elasticity, and freezing damage was accompanied by electrolyte leakage, consistent with cell-membrane disruption by ice. Both Chenopodiaceae and Poaceae had the potential to develop cold acclimation and withstand freezing during the growing season, which conflicted with the hypothesis. Instead, freezing tolerance was more closely associated with life history and ecological preference in these Mongolian species. PMID:23599273

Identification and characterization of miRNAs in two closely related C4 and C3 species of Cleome by high-throughput sequencing

PubMed Central

Gao, Shuangcheng; Zhao, Wei; Li, Xiang; You, Qingbo; Shen, Xinjie; Guo, Wei; Wang, Shihua; Shi, Guoan; Liu, Zheng; Jiao, Yongqing

2017-01-01

Cleome gynandra and Cleome hassleriana, which are C4 and C3 plants, respectively, are two species of Cleome. The close genetic relationship between C. gynandra and C. hassleriana provides advantages for discovering the differences in leaf development and physiological processes between C3 and C4 plants. MicroRNAs (miRNAs) are a class of important regulators of various biological processes. In this study, we investigate the differences in the characteristics of miRNAs between C. gynandra and C. hassleriana using high-throughput sequencing technology. In total, 94 and 102 known miRNAs were identified in C. gynandra and C. hassleriana, respectively, of which 3 were specific for C. gynandra and 10 were specific for C. hassleriana. Ninety-one common miRNAs were identified in both species. In addition, 4 novel miRNAs were detected, including three in C. gynandra and three in C. hassleriana. Of these miRNAs, 67 were significantly differentially expressed between these two species and were involved in extensive biological processes, such as glycol-metabolism and photosynthesis. Our study not only provided resources for C. gynandra and C. hassleriana research but also provided useful clues for the understanding of the roles of miRNAs in the alterations of biological processes in leaf tissues during the evolution of the C4 pathway. PMID:28422166

Arabidopsis ERG28 Tethers the Sterol C4-Demethylation Complex to Prevent Accumulation of a Biosynthetic Intermediate That Interferes with Polar Auxin Transport[C][W

PubMed Central

Mialoundama, Alexis Samba; Jadid, Nurul; Brunel, Julien; Di Pascoli, Thomas; Heintz, Dimitri; Erhardt, Mathieu; Mutterer, Jérôme; Bergdoll, Marc; Ayoub, Daniel; Van Dorsselaer, Alain; Rahier, Alain; Nkeng, Paul; Geoffroy, Philippe; Miesch, Michel; Camara, Bilal; Bouvier, Florence

2013-01-01

Sterols are vital for cellular functions and eukaryotic development because of their essential role as membrane constituents. Sterol biosynthetic intermediates (SBIs) represent a potential reservoir of signaling molecules in mammals and fungi, but little is known about their functions in plants. SBIs are derived from the sterol C4-demethylation enzyme complex that is tethered to the membrane by Ergosterol biosynthetic protein28 (ERG28). Here, using nonlethal loss-of-function strategies focused on Arabidopsis thaliana ERG28, we found that the previously undetected SBI 4-carboxy-4-methyl-24-methylenecycloartanol (CMMC) inhibits polar auxin transport (PAT), a key mechanism by which the phytohormone auxin regulates several aspects of plant growth, including development and responses to environmental factors. The induced accumulation of CMMC in Arabidopsis erg28 plants was associated with diagnostic hallmarks of altered PAT, including the differentiation of pin-like inflorescence, loss of apical dominance, leaf fusion, and reduced root growth. PAT inhibition by CMMC occurs in a brassinosteroid-independent manner. The data presented show that ERG28 is required for PAT in plants. Furthermore, it is accumulation of an atypical SBI that may act to negatively regulate PAT in plants. Hence, the sterol pathway offers further prospects for mining new target molecules that could regulate plant development. PMID:24326590

Initial Events during the Evolution of C4 Photosynthesis in C3 Species of Flaveria1[W][OPEN

PubMed Central

Sage, Tammy L.; Busch, Florian A.; Johnson, Daniel C.; Friesen, Patrick C.; Stinson, Corey R.; Stata, Matt; Sultmanis, Stefanie; Rahman, Beshar A.; Rawsthorne, Stephen; Sage, Rowan F.

2013-01-01

The evolution of C4 photosynthesis in many taxa involves the establishment of a two-celled photorespiratory CO2 pump, termed C2 photosynthesis. How C3 species evolved C2 metabolism is critical to understanding the initial phases of C4 plant evolution. To evaluate early events in C4 evolution, we compared leaf anatomy, ultrastructure, and gas-exchange responses of closely related C3 and C2 species of Flaveria, a model genus for C4 evolution. We hypothesized that Flaveria pringlei and Flaveria robusta, two C3 species that are most closely related to the C2 Flaveria species, would show rudimentary characteristics of C2 physiology. Compared with less-related C3 species, bundle sheath (BS) cells of F. pringlei and F. robusta had more mitochondria and chloroplasts, larger mitochondria, and proportionally more of these organelles located along the inner cell periphery. These patterns were similar, although generally less in magnitude, than those observed in the C2 species Flaveria angustifolia and Flaveria sonorensis. In F. pringlei and F. robusta, the CO2 compensation point of photosynthesis was slightly lower than in the less-related C3 species, indicating an increase in photosynthetic efficiency. This could occur because of enhanced refixation of photorespired CO2 by the centripetally positioned organelles in the BS cells. If the phylogenetic positions of F. pringlei and F. robusta reflect ancestral states, these results support a hypothesis that increased numbers of centripetally located organelles initiated a metabolic scavenging of photorespired CO2 within the BS. This could have facilitated the formation of a glycine shuttle between mesophyll and BS cells that characterizes C2 photosynthesis. PMID:24064930

Intermediate Temperature Strength Degradation in SiC/SiC Composites

NASA Technical Reports Server (NTRS)

Morscher, Gregory N.; Cawley, James D.; Levine, Stanley (Technical Monitor)

2001-01-01

Woven silicon carbide fiber-reinforced, silicon carbide matrix composites are leading candidate materials for an advanced jet engine combustor liner application. Although the use temperature in the hot region for this application is expected to exceed 1200 C, a potential life-limiting concern for this composite system exists at intermediate temperatures (800 +/- 200 C), where significant time-dependent strength degradation has been observed under stress-rupture loading. A number of factors control the degree of stress-rupture strength degradation, the major factor being the nature of the interphase separating the fiber and the matrix. BN interphases are superior to carbon interphases due to the slower oxidation kinetics of BN. A model for the intermediate temperature stress-rupture of SiC/BN/SiC composites is presented based on the observed mechanistic process that leads to strength degradation for the simple case of through-thickness matrix cracks. The approach taken has much in common with that used by Curtin and coworkers, for two different composite systems. The predictions of the model are in good agreement with the rupture data for stress-rupture of both precracked and as-produced composites. Also, three approaches that dramatically improve the intermediate temperature stress-rupture properties are described: Si-doped BN, fiber spreading, and 'outside debonding'.

Towards an integrative model of C4 photosynthetic subtypes: insights from comparative transcriptome analysis of NAD-ME, NADP-ME, and PEP-CK C4 species.

PubMed

Bräutigam, Andrea; Schliesky, Simon; Külahoglu, Canan; Osborne, Colin P; Weber, Andreas P M

2014-07-01

C4 photosynthesis affords higher photosynthetic carbon conversion efficiency than C3 photosynthesis and it therefore represents an attractive target for engineering efforts aiming to improve crop productivity. To this end, blueprints are required that reflect C4 metabolism as closely as possible. Such blueprints have been derived from comparative transcriptome analyses of C3 species with related C4 species belonging to the NAD-malic enzyme (NAD-ME) and NADP-ME subgroups of C4 photosynthesis. However, a comparison between C3 and the phosphoenolpyruvate carboxykinase (PEP-CK) subtype of C4 photosynthesis is still missing. An integrative analysis of all three C4 subtypes has also not been possible to date, since no comparison has been available for closely related C3 and PEP-CK C4 species. To generate the data, the guinea grass Megathyrsus maximus, which represents a PEP-CK species, was analysed in comparison with a closely related C3 sister species, Dichanthelium clandestinum, and with publicly available sets of RNA-Seq data from C4 species belonging to the NAD-ME and NADP-ME subgroups. The data indicate that the core C4 cycle of the PEP-CK grass M. maximus is quite similar to that of NAD-ME species with only a few exceptions, such as the subcellular location of transfer acid production and the degree and pattern of up-regulation of genes encoding C4 enzymes. One additional mitochondrial transporter protein was associated with the core cycle. The broad comparison identified sucrose and starch synthesis, as well as the prevention of leakage of C4 cycle intermediates to other metabolic pathways, as critical components of C4 metabolism. Estimation of intercellular transport fluxes indicated that flux between cells is increased by at least two orders of magnitude in C4 species compared with C3 species. In contrast to NAD-ME and NADP-ME species, the transcription of photosynthetic electron transfer proteins was unchanged in PEP-CK. In summary, the PEP-CK blueprint of M

Towards an integrative model of C4 photosynthetic subtypes: insights from comparative transcriptome analysis of NAD-ME, NADP-ME, and PEP-CK C4 species

PubMed Central

Bräutigam, Andrea; Schliesky, Simon; Külahoglu, Canan; Osborne, Colin P.; Weber, Andreas P.M.

2014-01-01

C4 photosynthesis affords higher photosynthetic carbon conversion efficiency than C3 photosynthesis and it therefore represents an attractive target for engineering efforts aiming to improve crop productivity. To this end, blueprints are required that reflect C4 metabolism as closely as possible. Such blueprints have been derived from comparative transcriptome analyses of C3 species with related C4 species belonging to the NAD-malic enzyme (NAD-ME) and NADP-ME subgroups of C4 photosynthesis. However, a comparison between C3 and the phosphoenolpyruvate carboxykinase (PEP-CK) subtype of C4 photosynthesis is still missing. An integrative analysis of all three C4 subtypes has also not been possible to date, since no comparison has been available for closely related C3 and PEP-CK C4 species. To generate the data, the guinea grass Megathyrsus maximus, which represents a PEP-CK species, was analysed in comparison with a closely related C3 sister species, Dichanthelium clandestinum, and with publicly available sets of RNA-Seq data from C4 species belonging to the NAD-ME and NADP-ME subgroups. The data indicate that the core C4 cycle of the PEP-CK grass M. maximus is quite similar to that of NAD-ME species with only a few exceptions, such as the subcellular location of transfer acid production and the degree and pattern of up-regulation of genes encoding C4 enzymes. One additional mitochondrial transporter protein was associated with the core cycle. The broad comparison identified sucrose and starch synthesis, as well as the prevention of leakage of C4 cycle intermediates to other metabolic pathways, as critical components of C4 metabolism. Estimation of intercellular transport fluxes indicated that flux between cells is increased by at least two orders of magnitude in C4 species compared with C3 species. In contrast to NAD-ME and NADP-ME species, the transcription of photosynthetic electron transfer proteins was unchanged in PEP-CK. In summary, the PEP-CK blueprint of M

Relative contribution of C3 and C4 type terrestrial organic matter in the Mahanadi offshore (Bay of Bengal) sediments and climatic implication.

NASA Astrophysics Data System (ADS)

da Silva, Rheane; Mazumdar, Aninda; Naik, Bg

2017-04-01

C3 and C4 are dominant vegetation in terrestrial environment. The primary product of photosynthesis of C3 plants is a 3 carbon bearing compound called phosphoglycerate (PGA). In contrast, CO2 is transferred to bundle sheath cells via 4 carbon bearing compound oxaloacetate/mallate and fixed by RuBiSCO in C4 plants. This marked variation in CO2 diffusion across stomata and enzymatic pathways lead to differences in stable carbon isotope ratios. Factors that control relative abundance of these vegetation types are concentration of p-CO2, temperature and humidity. Low p-CO2, air temperature below cross over temperature and aridity are the climatic parameters favoring expansion of C4 type vegetation, whereas higher extreme conditions promote greater C3 type production (Ehleringer, J. R, 2005). In marine sediment n-alkane (lipid fraction) distribution and compound specific isotope ratios are ideal markers to characterize nature of terrestrial organic flux owing to high diagenetic stability and near 100% extraction efficiency. We report here the relative abundance of C3-C4 vegetation over 8 marine isotope stages covering 300kyr. A 39.08 m long core (MD 161-19) was collected onboard ORV Marion Dufresne, at a water depth of 1480 m (Lat: 18 59.1092N Long: 85 41.1669E) (Mazumdar., et. al. 2014) for the study of sediment physico chemical properties and their link to paleoclimatic variation. The carbon isotope ratios of C-27 n-alkane range from -35.3‰ to -23.6‰ VPDB. 13C enrichment trends indicate a greater contribution from C4 vegetation type and 13C depletion trends are attributed to greater flux of C3 type vegetation. Mass balance calculation to reconstruct the temporal variation in C3/ C4 ratios is carried out using the end member values of -34.5‰ and -19.8‰ respectively (Collister.,et. al. 1994). The calculated C3/C4 ratio is 27:73 at LGM and shifts to 71:29 around 6 kyr BP. Based on results, we observe that colder isotope substages characterized by lower pCO2 saw

Pyrazolo[3,4-c]isothiazole and isothiazolo[4,3-d]isoxazole derivatives as antifungal agents.

PubMed

Vicentini, Chiara Beatrice; Romagnoli, Carlo; Manfredini, Stefano; Rossi, Damiano; Mares, Donatella

2011-05-01

The diseases of plants and humans due to pathogenic fungi are increasing. Among the substances used to combat fungi, the azoles are of primary interest, both in agricultural field both in health. To avoid fungal resistance phenomena, the synthesis and tests of new derivatives are necessary. This article discusses the synthesis and the antifungal activity of pyrazolo[3,4-c]isothiazole and isothiazolo[4,3-d]isoxazole derivatives against three fungi that are pathogenic only for plants and two fungi that are opportunistic in humans and plants. The compounds were prepared starting from 2-cyano-3-ethoxy-2-butenethioamide. The antifungal activity of the compounds was determined by measuring the inhibition of growth of the fungi tested at 20, 50, and 100 µg/mL in comparison with the controls. Results demonstrated that several compounds were able to control the mycelial growth of the tested fungi, even if they showed different sensitivity to the different azole-derivatives. In general Magnaporthe grisea (T.T. Hebert) Yaegashi & Udagawa was the most sensitive fungus, being blocked almost entirely by 4-chloro derivative even at 20 µg/mL, a concentration at which the reference commercial compound tricyclazole was nearly ineffective. These findings demonstrate that the pyrazolo[3,4-c]isothiazole derivatives have a wide spectrum of activity on phytopathogenic and opportunistic fungi. In particular the 4-chloro derivative seems to have a great potential as new product to combat M. grisea in the agricultural field.

NDVI, C3 and C4 production, and distributions in Great Plains grassland land cover classes

USGS Publications Warehouse

Tieszen, L.L.; Reed, Bradley C.; Bliss, Norman B.; Wylie, Bruce K.; DeJong, Benjamin D.

1997-01-01

The distributions of C3 and C4 grasses were used to interpret the distribution, seasonal performance, and potential production of grasslands in the Great Plains of North America. Thirteen major grassland seasonal land cover classes were studied with data from three distinct sources. Normalized Difference Vegetation Index (NDVI) data derived from the National Oceanic and Atmospheric Administration (NOAA) Advanced Very High Resolution Radiometer (AVHRR) sensor were collected for each pixel over a 5-yr period (1989–1993), analyzed for quantitative attributes and seasonal relationships, and then aggregated by land cover class. Data from the State Soil Geographic (STATSGO) database were used to identify dominant plant species contributing to the potential production in each map unit. These species were identified as C3 or C4, and contributions to production were aggregated to provide estimates of the percentage of C3 and C4 production for each intersection of the STATSGO map units and the seasonal land cover classes. Carbon isotope values were obtained at specific sites from the soil organic matter of the upper horizon of soil cores and were related to STATSGO estimates of potential production.The grassland classes were distributed with broad northwest-to-southeast orientations. Some classes had large variations in C3 and C4 composition with high proportions of C4species in the south and low proportions in the north. This diversity of photosynthetic types within land cover classes that cross regions of different temperature and precipitation results in similar seasonal patterns and magnitudes of NDVI. The easternmost class, 65, containing tallgrass prairie components, bluestem, Indiangrass, and switchgrass, possessed the highest maximum NDVI and time-integrated NDVI values each year. Grassland classes varied over 5 yr from a high integrated NDVI mean of 4.9 in class 65 in the east to a low of 1.2 in class 76 (sand sage, blue grama, wheatgrass, and buffalograss) in the

C4 photosynthesis and water stress

PubMed Central

Ghannoum, Oula

2009-01-01

Background In contrast to C3 photosynthesis, the response of C4 photosynthesis to water stress has been less-well studied in spite of the significant contribution of C4 plants to the global carbon budget and food security. The key feature of C4 photosynthesis is the operation of a CO2-concentrating mechanism in the leaves, which serves to saturate photosynthesis and suppress photorespiration in normal air. This article reviews the current state of understanding about the response of C4 photosynthesis to water stress, including the interaction with elevated CO2 concentration. Major gaps in our knowledge in this area are identified and further required research is suggested. Scope Evidence indicates that C4 photosynthesis is highly sensitive to water stress. With declining leaf water status, CO2 assimilation rate and stomatal conductance decrease rapidly and photosynthesis goes through three successive phases. The initial, mainly stomatal phase, may or may not be detected as a decline in assimilation rates depending on environmental conditions. This is because the CO2-concentrating mechanism is capable of saturating C4 photosynthesis under relatively low intercellular CO2 concentrations. In addition, photorespired CO2 is likely to be refixed before escaping the bundle sheath. This is followed by a mixed stomatal and non-stomatal phase and, finally, a mainly non-stomatal phase. The main non-stomatal factors include reduced activity of photosynthetic enzymes; inhibition of nitrate assimilation, induction of early senescence, and changes to the leaf anatomy and ultrastructure. Results from the literature about CO2 enrichment indicate that when C4 plants experience drought in their natural environment, elevated CO2 concentration alleviates the effect of water stress on plant productivity indirectly via improved soil moisture and plant water status as a result of decreased stomatal conductance and reduced leaf transpiration. Conclusions It is suggested that there is a

Generation of Anaphylatoxins by Human β-Tryptase from C3, C4, and C51

PubMed Central

Fukuoka, Yoshihiro; Xia, Han-Zhang; Sanchez-Muñoz, Laura B.; Dellinger, Anthony L.; Escribano, Luis; Schwartz, Lawrence B.

2009-01-01

Both mast cells and complement participate in innate and acquired immunity. The current study examines whether β-tryptase, the major protease of human mast cells, can directly generate bioactive complement anaphylatoxins. Important variables included pH, monomeric vs tetrameric forms of β-tryptase, and the β-tryptase-activating polyanion. The B12 mAb was used to stabilize β-tryptase in its monomeric form. C3a and C4a were best generated from C3 and C4, respectively, by monomeric β-tryptase in the presence of low molecular weight dextran sulfate or heparin at acidic pH. High molecular weight polyanions increased degradation of these anaphylatoxins. C5a was optimally generated from C5 at acidic pH by β-tryptase monomers in the presence of high molecular weight dextran sulfate and heparin polyanions, but also was produced by β-tryptase tetramers under these conditions. Mass spectrometry verified that the molecular mass of each anaphylatoxin was correct. Both β-tryptase-generated C5a and C3a (but not C4a) were potent activators of human skin mast cells. These complement anaphylatoxins also could be generated by β-tryptase in releasates of activated skin mast cells. Of further biologic interest, β-tryptase also generated C3a from C3 in human plasma at acidic pH. These results suggest β-tryptase might generate complement anaphylatoxins in vivo at sites of inflammation, such as the airway of active asthma patients where the pH is acidic and where elevated levels of β-tryptase and complement anaphylatoxins are detected. PMID:18424754

An rbcL mRNA-binding protein is associated with C3 to C4 evolution and light-induced production of Rubisco in Flaveria.

PubMed

Yerramsetty, Pradeep; Agar, Erin M; Yim, Won C; Cushman, John C; Berry, James O

2017-07-20

Nuclear-encoded RLSB protein binds chloroplastic rbcL mRNA encoding the Rubisco large subunit. RLSB is highly conserved across all groups of land plants and is associated with positive post-transcriptional regulation of rbcL expression. In C3 leaves, RLSB and Rubisco occur in all chlorenchyma cell chloroplasts, while in C4 leaves these accumulate only within bundle sheath (BS) chloroplasts. RLSB's role in rbcL expression makes modification of its localization a likely prerequisite for the evolutionary restriction of Rubisco to BS cells. Taking advantage of evolutionarily conserved RLSB orthologs in several C3, C3-C4, C4-like, and C4 photosynthetic types within the genus Flaveria, we show that low level RLSB sequence divergence and modification to BS specificity coincided with ontogeny of Rubisco specificity and Kranz anatomy during C3 to C4 evolution. In both C3 and C4 species, Rubisco production reflected RLSB production in all cell types, tissues, and conditions examined. Co-localization occurred only in photosynthetic tissues, and both proteins were co-ordinately induced by light at post-transcriptional levels. RLSB is currently the only mRNA-binding protein to be associated with rbcL gene regulation in any plant, with variations in sequence and acquisition of cell type specificity reflecting the progression of C4 evolution within the genus Flaveria. © The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.

Thermodynamics of triple helix formation: spectrophotometric studies on the d(A)10.2d(T)10 and d(C+3T4C+3).d(G3A4G3).d(C3T4C3) triple helices.

PubMed Central

Pilch, D S; Brousseau, R; Shafer, R H

1990-01-01

We have stabilized the d(A)10.2d(T)10 and d(C+LT4C+3).d(G3A4G3).d(C3T4C3) triple helices with either NaCl or MgCl2 at pH 5.5. UV mixing curves demonstrate a 1:2 stoichiometry of purine to pyrimidine strands under the appropriate conditions of pH and ionic strength. Circular dichroic titrations suggest a possible sequence-independent spectral signature for triplex formation. Thermal denaturation profiles indicate the initial loss of the third strand followed by dissociation of the underlying duplex with increasing temperature. Depending on the base sequence and ionic conditions, the binding affinity of the third strand for the duplex at 25 degrees C is two to five orders of magnitude lower than that of the two strands forming the duplex. Thermodynamic parameters for triplex formation were determined for both sequences in the presence of 50 mM MgCl2 and/or 2.0 M NaCl. Hoogsteen base pairs are 0.22-0.64 kcal/mole less stable than Watson-Crick base pairs, depending on ionic conditions and base composition. C+.G and T.A Hoogsteen base pairs appear to have similar stability in the presence of Mg2+ ions at low pH. PMID:2216768

Total Syntheses of (±)-Ovalicin, C4(S*)-Isomer, and Its C5-Analogs and Anti-trypanosomal Activities∥

PubMed Central

Hua, Duy H.; Zhao, Huiping; Battina, Srinivas K.; Lou, Kaiyan; Jimenez, Ana L.; Desper, John; Perchellet, Elisabeth M.; Perchellet, Jean-Pierre H.; Chiang, Peter K.

2008-01-01

Total syntheses of (±)-ovalicin, its C4(S*)-isomer 44, and C5-side chain intermediate 46 were accomplished via an intramolecular Heck reaction of (Z)-3-(t-butyldimethylsilyloxy)-1-iodo-1,6-heptadiene and a catalytic amount of palladium acetate. Subsequent epoxidation, dihydroxylation, methylation and oxidation led to (3S*,5R*,6R*)-5-methoxy-6-(t-butyldimethylsilyloxy)-1-oxaspiro[2.5]octan-4-one (2), a reported intermediate. The addition of a side chain with cis-1-lithio-1,5-dimethyl-1,4-hexadiene (27) followed by oxidation afforded (±)-ovalicin. The functional group manipulation afforded a number of regio- and stereoisomers, which allow the synthesis of analogs for bioevaluation. The structure of 44 was firmly established via a single-crystal X-ray analysis. The stereochemistry at C4 generated from the addition reactions of alkenyllithium with ketones 2, 40, and 45 is dictated by C6-alkoxy functionality. Anti-trypanosomal activities of various ovalicin analogs and synthetic intermediates were evaluated, and C5-side chain analog, 46, shows the strongest activity. Compound 44 shows antiproliferative effect against HL-60 tumor cells in vitro. Compounds 46 and a precursor, (3S*,4R*,5R*,6R*)-5-methoxy-4-[(E)-1’,5’-dimethylhexa-1’,4’–dienyl)]-6-(t-butyldimethylsilyloxy)-1-oxaspiro[2.5]octan-4-ol (28), may be explored for the development of anti-parasitic drugs. PMID:18356059

Allocate carbon for a reason: priorities are reflected in the ¹³C/¹²C ratios of plant lipids synthesized via three independent biosynthetic pathways.

PubMed

Zhou, Youping; Stuart-Williams, Hilary; Grice, Kliti; Kayler, Zachary E; Zavadlav, Saša; Vogts, Angela; Rommerskirchen, Florian; Farquhar, Graham D; Gessler, Arthur

2015-03-01

It has long been theorized that carbon allocation, in addition to the carbon source and to kinetic isotopic effects associated with a particular lipid biosynthetic pathway, plays an important role in shaping the carbon isotopic composition ((13)C/(12)C) of lipids (Park and Epstein, 1961). If the latter two factors are properly constrained, valuable information about carbon allocation during lipid biosynthesis can be obtained from carbon isotope measurements. Published work of Chikaraishi et al. (2004) showed that leaf lipids isotopic shifts from bulk leaf tissue Δδ(13)C(bk-lp) (defined as δ(13)C(bulkleaftissue)-δ(13)C(lipid)) are pathway dependent: the acetogenic (ACT) pathway synthesizing fatty lipids has the largest isotopic shift, the mevalonic acid (MVA) pathway synthesizing sterols the lowest and the phytol synthesizing 1-deoxy-D-xylulose 5-phosphate (DXP) pathway gives intermediate values. The differences in Δδ(13)C(bk-lp) between C3 and C4 plants Δδ(13)C(bk-lp,C4-C3) are also pathway-dependent: Δδ(13)C(ACT)(bk-lp,C4-C3) > Δδ(13)C(DXP(bk-lp,C4-C3) > Δδ(13)C(MVA)(bk-lp,C4-C3). These pathway-dependent differences have been interpreted as resulting from kinetic isotopic effect differences of key but unspecified biochemical reactions involved in lipids biosynthesis between C3 and C4 plants. After quantitatively considering isotopic shifts caused by (dark) respiration, export-of-carbon (to sink tissues) and photorespiration, we propose that the pathway-specific differences Δδ(13)C(bk-lp,C4-C3) can be successfully explained by C4-C3 carbon allocation (flux) differences with greatest flux into the ACT pathway and lowest into the MVA pathways (when flux is higher, isotopic shift relative to source is smaller). Highest carbon allocation to the ACT pathway appears to be tied to the most stringent role of water-loss-minimization by leaf waxes (composed mainly of fatty lipids) while the lowest carbon allocation to the MVA pathway can be largely explained

Kranz and single-cell forms of C4 plants in the subfamily Suaedoideae show kinetic C4 convergence for PEPC and Rubisco with divergent amino acid substitutions

PubMed Central

Rosnow, Josh J.; Evans, Marc A.; Kapralov, Maxim V.; Cousins, Asaph B.; Edwards, Gerald E.; Roalson, Eric H.

2015-01-01

The two carboxylation reactions performed by phosphoenolpyruvate carboxylase (PEPC) and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) are vital in the fixation of inorganic carbon for C4 plants. The abundance of PEPC is substantially elevated in C4 leaves, while the location of Rubisco is restricted to one of two chloroplast types. These differences compared with C3 leaves have been shown to result in convergent enzyme optimization in some C4 species. Investigation into the kinetic properties of PEPC and Rubisco from Kranz C4, single cell C4, and C3 species in Chenopodiaceae s. s. subfamily Suaedoideae showed that these major carboxylases in C4 Suaedoideae species lack the same mutations found in other C4 systems which have been examined; but still have similar convergent kinetic properties. Positive selection analysis on the N-terminus of PEPC identified residues 364 and 368 to be under positive selection with a posterior probability >0.99 using Bayes empirical Bayes. Compared with previous analyses on other C4 species, PEPC from C4 Suaedoideae species have different convergent amino acids that result in a higher K m for PEP and malate tolerance compared with C3 species. Kinetic analysis of Rubisco showed that C4 species have a higher catalytic efficiency of Rubisco (k catc in mol CO2 mol–1 Rubisco active sites s–1), despite lacking convergent substitutions in the rbcL gene. The importance of kinetic changes to the two-carboxylation reactions in C4 leaves related to amino acid selection is discussed. PMID:26417023

The electron affinities of C{sub 3}O and C{sub 4}O

DOE Office of Scientific and Technical Information (OSTI.GOV)

Rienstra-Kiracofe, J.C.; Ellison, G.B.; Hoffman, B.C.

The authors predict the adiabatic electron affinities of C{sub 3}O and C{sub 4}O based on electronic structure calculations, using a large triple-{zeta} basis set with polarization and diffuse functions (TZ2Pf+diff) with the SCF, CCSD, and CCSD(T) methods as well as with the aug-cc-pVDZ and aug-cc-pVTZ basis sets. The results imply electron affinities for C{sub 3}O and C{sub 4}O; EA(C{sub 3}O) = 0.93 eV {+-} 0.10 and EA(C{sub 4}O) = 2.99 {+-} 0.10. The EA(C{sub 3}O) is 0.41 eV lower than the experimental value of 1.34 {+-} 0.15 eV, while the EA(C{sub 4}O) is 0.94 eV higher than the experimental valuemore » of 2.05 {+-} 0.15 eV. Optimized geometries for all species at each level of theory are given, and harmonic vibrational frequencies are reported at the SCF/TZ2Pf+diff and CCSD/aug-cc-pVDZ levels.« less

Investigations of electron attachment to the perfluorocarbon molecules c-C4F8, 2-C4F8, 1,3 C4F6, and c-C5F8

NASA Astrophysics Data System (ADS)

Feil, Stefan; Märk, Tilmann D.; Mauracher, Andreas; Scheier, Paul; Mayhew, Chris A.

2008-11-01

Non-dissociative and dissociative electron attachment to a series of gas-phase perfluorocarbons (PFCs), namely octafluorocyclobutane, c-C4F8, octafluorobut-2-ene (perfluoro-2-butene), 2-C4F8, hexafluorobuta-1,3-diene (1,3 perfluorobutadiene), 1,3 C4F6, and octafluorocyclopentene (perfluorocyclopentene), c-C5F8, of importance to technological plasmas, have been investigated using two different, but complimentary, instruments available in Innsbruck over the electron energy range 0-20 eV. Anion yields as a function of electron energy have been recorded, with the positions and intensities of the electron attachment resonances being determined. One of these instruments is a double focusing sector field mass spectrometer (VG-ZAB-2SEQ), which has been used for measurements requiring high sensitivity and for obtaining accurate relative anion yields. It has also been used to determine the electron detachment lifetimes of the parent anions under various accelerating voltages, and these results are also presented. The second instrument (CELIA) is a trochoidal electron monochromator coupled to a quadrupole mass filter with a pulse counting system for detecting product anionic species. This provides a much higher energy resolution than the VG-ZAB, which makes it a better instrument to investigate narrow energy resonances close to 0 eV. The results of anion yields, peak positions and the relative intensities presented in this paper are compared with previous data of electron attachment to the above PFCs, including investigations by Professor Eugen Illenberger.

Cellular expression of C3 and C4 photosynthetic enzymes in the amphibious sedge Eleocharis retroflexa ssp. chaetaria.

PubMed

Ueno, Osamu; Wakayama, Masataka

2004-12-01

The amphibious leafless sedge Eleocharis retroflexa ssp. chaetaria expresses C(4)-like biochemical characteristics in both the terrestrial and submerged forms. Culms of the terrestrial form have Kranz anatomy, whereas those of the submerged form have Kranz-like anatomy combined with anatomical features of aquatic plant leaves. We examined the immunolocalization of C(3) and C(4) enzymes in culms of the two forms. In both forms, phosphoenolpyruvate carboxylase; pyruvate, Pi dikinase; and NAD-malic enzyme were compartmentalized between the mesophyll (M) and Kranz cells, but their levels were somewhat reduced in the submerged form. In the terrestrial form, ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco) occurred mainly in the Kranz cells, and weakly in the M chloroplasts. In the submerged form, the rubisco occurred at higher levels in the M cells than in the terrestrial form. In both forms, the C(4) pattern of enzyme expression was clearer in the M cells adjacent to Kranz cells than in distant M cells. During the transition from terrestrial to submerged conditions, the enzyme expression pattern changed in submerged mature culms that had been formed in air before submergence, and matched that in culms newly developed underwater. It seems that effects of both environmental and developmental factors overlap in the C(4) pattern expression in this plant.

Water-assisted production of honeycomb-like g-C3N4 with ultralong carrier lifetime and outstanding photocatalytic activity

NASA Astrophysics Data System (ADS)

Wang, Zhenyu; Guan, Wei; Sun, Yanjuan; Dong, Fan; Zhou, Ying; Ho, Wing-Kei

2015-01-01

Graphitic carbon nitride (g-C3N4) is a visible light photocatalyst, limited by low activity mainly caused by rapid recombination of charge carriers. In the present work, honeycomb-like g-C3N4 was synthesized via thermal condensation of urea with addition of water at 450 °C for 1 h. Prolonging the condensation time caused the morphology of g-C3N4 to change from a porous honeycomb structure to a velvet-like nanoarchitecture. Unlike in previous studies, the photocatalytic activity of g-C3N4 decreased with increasing surface area. The honeycomb-like g-C3N4 with a relatively low surface area showed highly enhanced photocatalytic activity with an NO removal ratio of 48%. The evolution of NO2 intermediate was dramatically inhibited over the honeycomb-like g-C3N4. The short and long lifetimes of the charge carriers for honeycomb-like g-C3N4 were unprecedentedly prolonged to 22.3 and 165.4 ns, respectively. As a result, the honeycomb-like g-C3N4 was highly efficient and stable in activity and could be used repeatedly. Addition of water had the following multiple positive effects on g-C3N4: (1) formation of the honeycomb structure, (2) promotion of charge separation and migration, (3) enlargement of the band gap, (4) increase in production yield, and (5) decrease in energy cost. These advantages make the present preparation method for highly efficient g-C3N4 extremely appealing for large-scale applications. The active species produced from g-C3N4 under illumination were confirmed using DMPO-ESR spin-trapping, the reaction intermediate was monitored, and the reaction mechanism of photocatalytic NO oxidation by g-C3N4 was revealed. This work could provide an attractive alternative method for mass-production of highly active g-C3N4-based photocatalysts for environmental and energetic applications.Graphitic carbon nitride (g-C3N4) is a visible light photocatalyst, limited by low activity mainly caused by rapid recombination of charge carriers. In the present work, honeycomb

New evidence for grain specific C4 photosynthesis in wheat

PubMed Central

Rangan, Parimalan; Furtado, Agnelo; Henry, Robert J

2016-01-01

The C4 photosynthetic pathway evolved to allow efficient CO2 capture by plants where effective carbon supply may be limiting as in hot or dry environments, explaining the high growth rates of C4 plants such as maize. Important crops such as wheat and rice are C3 plants resulting in efforts to engineer them to use the C4 pathway. Here we show the presence of a C4 photosynthetic pathway in the developing wheat grain that is absent in the leaves. Genes specific for C4 photosynthesis were identified in the wheat genome and found to be preferentially expressed in the photosynthetic pericarp tissue (cross- and tube-cell layers) of the wheat caryopsis. The chloroplasts exhibit dimorphism that corresponds to chloroplasts of mesophyll- and bundle sheath-cells in leaves of classical C4 plants. Breeding to optimize the relative contributions of C3 and C4 photosynthesis may adapt wheat to climate change, contributing to wheat food security. PMID:27530078

Leaf and plant water use efficiency of C{sub 4} species grown at glacial to elevated CO{sub 2} concentrations

DOE Office of Scientific and Technical Information (OSTI.GOV)

Polley, H.W.; Johnson, H.B.; Mayeux, H.S.

1996-03-01

Leaf gas exchange was measured on C{sub 4} plants grown from near glacial to current CO{sub 2} concentrations (200-350 {mu}mol mol{sup -1}) and from the current concentration to possible future levels (near 700 and 1000 {mu}mol mol{sup -1}) to test the prediction that intrinsic water use efficiency (CO{sub 2} assimilation [A]/stomatal conductance to water [g]) would rise by a similar relative amount as CO{sub 2} concentration. Studied were species differing in growth form or life history, the perennial grass Schizachyrium scoparium (little bluestem), perennial shrub Atriplex canescens (four-wing saltbush), and annual grass Schizachyrium scoparium (little bluestem), leaf A/g of themore » C{sub 4} species examined was stimulated proportionally more by a given relative increase in CO{sub 2} over subambient than by elevated concentrations. The ratio of the relative increase in A/g to that in CO{sub 2} exceeded unity in S, scoparium and A. canescens as CO{sub 2} rose from 700 to 1000 {mu}mol mol{sup -1}. At higher CO{sub 2} concentrations, A/g of the C{sub 4} perennials was similar to that expected for C{sub 3} plants. Since much of the potential response of C{sub 4} plants to CO{sub 4} perennials was similar to that expected for C{sub 3} plants. Since much of the potential response of C{sub 4} plants to CO{sub 2} often derives from higher water use efficiency (WUE), these results indicated that potential productivity of some C{sub 4} plants increased relatively more since glaciation than it will in the future. There also were large (>100%) differences in A/g and plant WUE (production/transpiration) at a given CO{sub 2} level among the plants examined that could influence the relative productivities of C{sub 4} species or growth forms and their interactions with C{sub 3} plants. 34 refs., 3 figs., 3 tabs.« less

Promotion of Cyclic Electron Transport Around Photosystem I with the Development of C4 Photosynthesis.

PubMed

Munekage, Yuri Nakajima; Taniguchi, Yukimi Y

2016-05-01

C4 photosynthesis is present in approximately 7,500 species classified into 19 families, including monocots and eudicots. In the majority of documented cases, a two-celled CO2-concentrating system that uses a metabolic cycle of four-carbon compounds is employed. C4 photosynthesis repeatedly evolved from C3 photosynthesis, possibly driven by the survival advantages it bestows in the hot, often dry, and nutrient-poor soils of the tropics and subtropics. The development of the C4 metabolic cycle greatly increased the ATP demand in chloroplasts during the evolution of malic enzyme-type C4 photosynthesis, and the additional ATP required for C4 metabolism may be produced by the cyclic electron transport around PSI. Recent studies have revealed the nature of cyclic electron transport and the elevation of its components during C4 evolution. In this review, we discuss the energy requirements of C3 and C4 photosynthesis, the current model of cyclic electron transport around PSI and how cyclic electron transport is promoted during C4 evolution using studies on the genus Flaveria, which contains a number of closely related C3, C4 and C3-C4 intermediate species. © The Author 2016. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.

Malate decarboxylases: evolution and roles of NAD(P)-ME isoforms in species performing C(4) and C(3) photosynthesis.

PubMed

Maier, Alexandra; Zell, Martina B; Maurino, Veronica G

2011-05-01

In the C(4) pathway of photosynthesis two types of malate decarboxylases release CO(2) in bundle sheath cells, NADP- and NAD-dependent malic enzyme (NADP-ME and NAD-ME), located in the chloroplasts and the mitochondria of these cells, respectively. The C(4) decarboxylases involved in C(4) photosynthesis did not evolve de novo; they were recruited from existing housekeeping isoforms. NADP-ME housekeeping isoforms would function in the control of malate levels during hypoxia, pathogen defence responses, and microspore separation, while NAD-ME participates in the respiration of malate in the tricarboxylic acid cycle. Recently, the existence of three enzymatic NAD-ME entities in Arabidopsis, occurring by alternative association of two subunits, was described as a novel mechanism to regulate NAD-ME activity under changing metabolic environments. The C(4) NADP-ME is thought to have evolved from a C(3) chloroplastic ancestor, which in turn would have evolved from an ancient cytosolic enzyme. In this way, the C(4) NADP-ME would have emerged through gene duplication, acquisition of a new promoter, and neo-functionalization. In contrast, there would exist a unique NAD-ME in C(4) plants, which would have been adapted to perform a dual function through changes in the kinetic and regulatory properties of the C(3) ancestors. In addition to this, for the evolution of C(4) NAD-ME, insertion of promoters or enhancers into the single-copy genes of the C(3) ancestors would have changed the expression without gene duplication.

Kranz and single-cell forms of C4 plants in the subfamily Suaedoideae show kinetic C4 convergence for PEPC and Rubisco with divergent amino acid substitutions.

PubMed

Rosnow, Josh J; Evans, Marc A; Kapralov, Maxim V; Cousins, Asaph B; Edwards, Gerald E; Roalson, Eric H

2015-12-01

The two carboxylation reactions performed by phosphoenolpyruvate carboxylase (PEPC) and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) are vital in the fixation of inorganic carbon for C4 plants. The abundance of PEPC is substantially elevated in C4 leaves, while the location of Rubisco is restricted to one of two chloroplast types. These differences compared with C3 leaves have been shown to result in convergent enzyme optimization in some C4 species. Investigation into the kinetic properties of PEPC and Rubisco from Kranz C4, single cell C4, and C3 species in Chenopodiaceae s. s. subfamily Suaedoideae showed that these major carboxylases in C4 Suaedoideae species lack the same mutations found in other C4 systems which have been examined; but still have similar convergent kinetic properties. Positive selection analysis on the N-terminus of PEPC identified residues 364 and 368 to be under positive selection with a posterior probability >0.99 using Bayes empirical Bayes. Compared with previous analyses on other C4 species, PEPC from C4 Suaedoideae species have different convergent amino acids that result in a higher K m for PEP and malate tolerance compared with C3 species. Kinetic analysis of Rubisco showed that C4 species have a higher catalytic efficiency of Rubisco (k catc in mol CO2 mol(-1) Rubisco active sites s(-1)), despite lacking convergent substitutions in the rbcL gene. The importance of kinetic changes to the two-carboxylation reactions in C4 leaves related to amino acid selection is discussed. © The Author 2015. Published by Oxford University Press on behalf of the Society for Experimental Biology.

Spatial distributions ofC3 and C4 grass functional types in the U.S. great plains and their despendency on inter-annual climate variability

USDA-ARS?s Scientific Manuscript database

Grassland ecosystems in North America are primarily composed of C3 and C4 plant functional types (PFTs) with their relative cover varying spatially and temporally. This study used 500-m MODIS surface reflectance products (MOD09A1) from 2000 to 2009 to extract an NDVI time series of C3 and C4 PFTs in...

X-ray diffraction analysis of 4- and 4'-substituted C n H2 n + 1O-C6H3(OH)-CH=N-C6H4-C m H2 m + 1 ( n/ m = 2/1 and 3/4) salicylideneanilines

NASA Astrophysics Data System (ADS)

Kuz'mina, L. G.; Navasardyan, M. A.; Mikhailov, A. A.

2017-11-01

X-ray diffraction study of two crystalline modifications of C2H5O-C6H3(OH)-CH=N-C6H4-CH3 ( 1a, sp. gr. P21/ n, and 1b, sp. gr. C2/c) and C3H7O-C6H3(OH)-CH=N-C6H4-C4H9 ( 2, sp. gr. P212121) has been performed. The 1a crystal structure contains two independent molecules. The molecules are conformationally nonrigid with respect to the mutual rotation of benzene rings; the dihedral angles between their planes are 29.19° and 26.00° in the independent molecules of 1a, 18.72° in the molecule of 1b, and 50.35° in the molecule of 2. The crystal packing of the compounds is discussed.

Characterization of a n+3C/n−4H SiC heterojunction diode

DOE Office of Scientific and Technical Information (OSTI.GOV)

Minamisawa, R. A.; Mihaila, A.; Farkas, I.

We report on the fabrication of n + 3C/n-4H SiC heterojunction diodes (HJDs) potentially promising the ultimate thermal stability of the junction. The diodes were systematically analyzed by TEM, X-ray diffraction, AFM, and secondary ion mass spectroscopy, indicating the formation of epitaxial 3C-SiC crystal on top of 4H-SiC substrate with continuous interface, low surface roughness, and up to ∼7 × 10{sup 17 }cm{sup −3} dopant impurity concentration. The conduction band off-set is about 1 V as extracted from CV measurements, while the valence bands of both SiC polytypes are aligned. The HJDs feature opening voltage of 1.65 V, consistent with the barrier height of about 1.5 eV extractedmore » from CV measurement. We finally compare the electrical results of the n + 3C/n-4H SiC heterojunction diodes with those featuring Si and Ge doped anodes in order to evaluate current challenges involved in the fabrication of such devices.« less

Improving our understanding of environmental controls on the distribution of C3 and C4 grasses.

PubMed

Pau, Stephanie; Edwards, Erika J; Still, Christopher J

2013-01-01

A number of studies have demonstrated the ecological sorting of C3 and C4 grasses along temperature and moisture gradients. However, previous studies of C3 and C4 grass biogeography have often inadvertently compared species in different and relatively unrelated lineages, which are associated with different environmental settings and distinct adaptive traits. Such confounded comparisons of C3 and C4 grasses may bias our understanding of ecological sorting imposed strictly by photosynthetic pathway. Here, we used MaxEnt species distribution modeling in combination with satellite data to understand the functional diversity of C3 and C4 grasses by comparing both large clades and closely related sister taxa. Similar to previous work, we found that C4 grasses showed a preference for regions with higher temperatures and lower precipitation compared with grasses using the C3 pathway. However, air temperature differences were smaller (2 °C vs. 4 °C) and precipitation and % tree cover differences were larger (1783 mm vs. 755 mm, 21.3% vs. 7.7%, respectively) when comparing C3 and C4 grasses within the same clade vs. comparing all C4 and all C3 grasses (i.e., ignoring phylogenetic structure). These results were due to important differences in the environmental preferences of C3 BEP and PACMAD clades (the two main grass clades). Winter precipitation was found to be more important for understanding the distribution and environmental niche of C3 PACMADs in comparison with both C3 BEPs and C4 taxa, for which temperature was much more important. Results comparing closely related C3 -C4 sister taxa supported the patterns derived from our modeling of the larger clade groupings. Our findings, which are novel in comparing the distribution and niches of clades, demonstrate that the evolutionary history of taxa is important for understanding the functional diversity of C3 and C4 grasses, and should have implications for how grasslands will respond to global change. © 2012

Laboratory detection of the C3N an C4H free radicals

NASA Technical Reports Server (NTRS)

Gottlieb, C. A.; Gottlieb, E. W.; Thaddeus, P.; Kawamura, H.

1983-01-01

The millimeter-wave spectra of the linear carbon chain free radicals C3N and C4H, first identified in IRC + 10216 and hitherto observed only in a few astronomical sources, have been detected with a Zeeman-modulated spectrometer in laboratory glow discharges through low pressure flowing mixtures of N2 + HC3N and He + HCCH, respectively. Four successive rotational transitions between 168 and 198 GHz have been measured for C3N, and five rotational transitions between 143 and 200 GHz for C4H; each is a well-resolved spin doublet owing to the unpaired electron present in both species. Precise values for the rotational, centrifugal distortion, and spin doubling constants have been obtained, which, with hyperfine constants derived from observations of the lower rotational transitions in the astronomical source TMC 1, allow all the rotational transitions of C3N and C4H at frequencies less than 300 GHz to be calculated to an absolute accuracy exceeding 1 ppm.

Water use patterns of co-occurring C3 and C4 shrubs in the Gurbantonggut desert in northwestern China.

PubMed

Tiemuerbieke, Bahejiayinaer; Min, Xiao-Jun; Zang, Yong-Xin; Xing, Peng; Ma, Jian-Ying; Sun, Wei

2018-09-01

In water-limited ecosystems, spatial and temporal partitioning of water sources is an important mechanism that facilitates plant survival and lessens the competition intensity of co-existing plants. Insights into species-specific root functional plasticity and differences in the water sources of co-existing plants under changing water conditions can aid in accurate prediction of the response of desert ecosystems to future climate change. We used stable isotopes of soil water, groundwater and xylem water to determine the seasonal and inter- and intraspecific differences variations in the water sources of six C 3 and C 4 shrubs in the Gurbantonggut desert. We also measured the stem water potentials to determine the water stress levels of each species under varying water conditions. The studied shrubs exhibited similar seasonal water uptake patterns, i.e., all shrubs extracted shallow soil water recharged by snowmelt water during early spring and reverted to deeper water sources during dry summer periods, indicating that all of the studied shrubs have dimorphic root systems that enable them to obtain water sources that differ in space and time. Species in the C 4 shrub community exhibited differences in seasonal water absorption and water status due to differences in topography and rooting depth, demonstrating divergent adaptations to water availability and water stress. Haloxylon ammodendron and T. ramosissima in the C 3 /C 4 mixed community were similar in terms of seasonal water extraction but differed with respect to water potential, which indicated that plant water status is controlled by both root functioning and shoot eco-physiological traits. The two Tamarix species in the C 3 shrub community were similar in terms of water uptake and water status, which suggests functional convergence of the root system and physiological performance under same soil water conditions. In different communities, Haloxylon ammodendron differed in terms of summer water extraction

Fabrication of modified g-C3N4 nanorod/Ag3PO4 nanocomposites for solar-driven photocatalytic oxygen evolution from water splitting

NASA Astrophysics Data System (ADS)

Tian, Lin; Xian, Xiaozhai; Cui, Xingkai; Tang, Hua; Yang, Xiaofei

2018-02-01

Semiconductor-based photocatalysis has been considered as one of the most effective techniques to achieve the conversion of clean and sustainable sunlight to solar fuel, in which the construction of novel solar-driven photocatalytic systems is the key point. Here, we report initially the synthesis of modified graphitic carbon nitride (g-C3N4) nanorods via the calcination of intermediates obtained from the co-polymerization of precursors, and the in-situ hybridization of Ag3PO4 with as-prepared modified g-C3N4 to produce g-C3N4 nanorod/Ag3PO4 composite materials. The diameter of modified rod-like g-C3N4 materials is determined to be around 1 μm. Subsequently the morphological features, crystal and chemical structures of the assembled g-C3N4 nanorod/Ag3PO4 composites were systematically investigated by SEM, XRD, XPS, UV-vis diffuse reflectance spectra (DRS). Furthermore, the use of as-prepared composite materials as the catalyst for photocatalytic oxygen evolution from water splitting was studied. The oxygen-generating results showed that the composite photocatalyst modified with 600 mg rod-like g-C3N4 demonstrates 2.5 times higher efficiency than that of bulk Ag3PO4. The mechanism behind the enhancement in the oxygen-evolving activity is proposed on the basis of in-situ electron spin resonance (ESR) measurement as well as theoretical analysis. The study provides new insights into the design and development of new photocatalytic composite materials for energy and environmental applications.

Stable Carbon Isotope Ratios of Individual Pollen Grains as a Proxy for C3- Versus C4-Grass Abundance in Paleorecords: A Validation Study

NASA Astrophysics Data System (ADS)

Nelson, D. M.; Hu, F.; Pearson, A.

2007-12-01

distinguishing C3 and C4 shifts. At 9 of the 10 sites C/N values indicate that surface-sediment organic matter was derived primarily from aquatic production. At the one site where organic matter was produced primarily by vascular plants the δ13C value (-29.3°) suggests organic matter derived exclusively from C3 plants. However, ~80% of the grasses on the landscape at this site are C4 grasses. The C3- like bulk-sediment δ13C value likely represents woody species, which comprise >90% of the pollen spectra. Thus δ13C analysis of single grains of grass pollen offers a new tool to classify grass pollen into two major functional groups and promises to advance our understanding of grassland ecology and evolution. Reference Nelson, D.M., Hu, F.S., Mikucki, J., Tian, J., and Pearson, A., 2007, Carbon isotopic analysis of individual pollen grains from C3 and C4 grasses using a spooling wire microcombustion interface: Geochimica et Cosmochimica Acta, v. 71, p. 4005-4014.

Intron loss from the NADH dehydrogenase subunit 4 gene of lettuce mitochondrial DNA: evidence for homologous recombination of a cDNA intermediate.

PubMed

Geiss, K T; Abbas, G M; Makaroff, C A

1994-04-01

The mitochondrial gene coding for subunit 4 of the NADH dehydrogenase complex I (nad4) has been isolated and characterized from lettuce, Lactuca sativa. Analysis of nad4 genes in a number of plants by Southern hybridization had previously suggested that the intron content varied between species. Characterization of the lettuce gene confirms this observation. Lettuce nad4 contains two exons and one group IIA intron, whereas previously sequenced nad4 genes from turnip and wheat contain three group IIA introns. Northern analysis identified a transcript of 1600 nucleotides, which represents the mature nad4 mRNA and a primary transcript of 3200 nucleotides. Sequence analysis of lettuce and turnip nad4 cDNAs was used to confirm the intron/exon border sequences and to examine RNA editing patterns. Editing is observed at the 5' and 3' ends of the lettuce transcript, but is absent from sequences that correspond to exons two, three and the 5' end of exon four in turnip and wheat. In contrast, turnip transcripts are highly edited in this region, suggesting that homologous recombination of an edited and spliced cDNA intermediate was involved in the loss of introns two and three from an ancestral lettuce nad4 gene.

Setaria viridis: A Model for C4 Photosynthesis[C][W

PubMed Central

Brutnell, Thomas P.; Wang, Lin; Swartwood, Kerry; Goldschmidt, Alexander; Jackson, David; Zhu, Xin-Guang; Kellogg, Elizabeth; Van Eck, Joyce

2010-01-01

C4 photosynthesis drives productivity in several major food crops and bioenergy grasses, including maize (Zea mays), sugarcane (Saccharum officinarum), sorghum (Sorghum bicolor), Miscanthus x giganteus, and switchgrass (Panicum virgatum). Gains in productivity associated with C4 photosynthesis include improved water and nitrogen use efficiencies. Thus, engineering C4 traits into C3 crops is an attractive target for crop improvement. However, the lack of a small, rapid cycling genetic model system to study C4 photosynthesis has limited progress in dissecting the regulatory networks underlying the C4 syndrome. Setaria viridis is a member of the Panicoideae clade and is a close relative of several major feed, fuel, and bioenergy grasses. It is a true diploid with a relatively small genome of ~510 Mb. Its short stature, simple growth requirements, and rapid life cycle will greatly facilitate genetic studies of the C4 grasses. Importantly, S. viridis uses an NADP-malic enzyme subtype C4 photosynthetic system to fix carbon and therefore is a potentially powerful model system for dissecting C4 photosynthesis. Here, we summarize some of the recent advances that promise greatly to accelerate the use of S. viridis as a genetic system. These include our recent successful efforts at regenerating plants from seed callus, establishing a transient transformation system, and developing stable transformation. PMID:20693355

Loss of the Chloroplast Transit Peptide from an Ancestral C3 Carbonic Anhydrase Is Associated with C4 Evolution in the Grass Genus Neurachne1[OPEN

PubMed Central

Clayton, Harmony; Saladié, Montserrat; Sharwood, Robert; Macfarlane, Terry

2017-01-01

Neurachne is the only known grass lineage containing closely related C3, C3-C4 intermediate, and C4 species, making it an ideal taxon with which to study the evolution of C4 photosynthesis in the grasses. To begin dissecting the molecular changes that led to the evolution of C4 photosynthesis in this group, the complementary DNAs encoding four distinct β-carbonic anhydrase (CA) isoforms were characterized from leaf tissue of Neurachne munroi (C4), Neurachne minor (C3-C4), and Neurachne alopecuroidea (C3). Two genes (CA1 and CA2) each encode two different isoforms: CA1a/CA1b and CA2a/CA2b. Transcript analyses found that CA1 messenger RNAs were significantly more abundant than transcripts from the CA2 gene in the leaves of each species examined, constituting ∼99% of all β-CA transcripts measured. Localization experiments using green fluorescent protein fusion constructs showed that, while CA1b is a cytosolic CA in all three species, the CA1a proteins are differentially localized. The N. alopecuroidea and N. minor CA1a isoforms were imported into chloroplasts of Nicotiana benthamiana leaf cells, whereas N. munroi CA1a localized to the cytosol. Sequence analysis indicated an 11-amino acid deletion in the amino terminus of N. munroi CA1a relative to the C3 and C3-C4 proteins, suggesting that chloroplast targeting of CA1a is the ancestral state and that loss of a functional chloroplast transit peptide in N. munroi CA1a is associated with the evolution of C4 photosynthesis in Neurachne spp. Remarkably, this mechanism is homoplastic with the evolution of the C4-associated CA in the dicotyledonous genus Flaveria, although the actual mutations in the two lineages differ. PMID:28153918

Evolution of C4 plants: a new hypothesis for an interaction of CO2 and water relations mediated by plant hydraulics.

PubMed

Osborne, Colin P; Sack, Lawren

2012-02-19

C(4) photosynthesis has evolved more than 60 times as a carbon-concentrating mechanism to augment the ancestral C(3) photosynthetic pathway. The rate and the efficiency of photosynthesis are greater in the C(4) than C(3) type under atmospheric CO(2) depletion, high light and temperature, suggesting these factors as important selective agents. This hypothesis is consistent with comparative analyses of grasses, which indicate repeated evolutionary transitions from shaded forest to open habitats. However, such environmental transitions also impact strongly on plant-water relations. We hypothesize that excessive demand for water transport associated with low CO(2), high light and temperature would have selected for C(4) photosynthesis not only to increase the efficiency and rate of photosynthesis, but also as a water-conserving mechanism. Our proposal is supported by evidence from the literature and physiological models. The C(4) pathway allows high rates of photosynthesis at low stomatal conductance, even given low atmospheric CO(2). The resultant decrease in transpiration protects the hydraulic system, allowing stomata to remain open and photosynthesis to be sustained for longer under drying atmospheric and soil conditions. The evolution of C(4) photosynthesis therefore simultaneously improved plant carbon and water relations, conferring strong benefits as atmospheric CO(2) declined and ecological demand for water rose.

Bundle-sheath thylakoids from NADP-malic enzyme-type C4 plants require an exogenous electron donor for enzyme light activation.

PubMed

Lavergne, D; Droux, M; Jacquot, J P; Miginiac-Maslow, M; Champigny, M L; Gadal, P

1985-10-01

Light activation of either NADP-malate dehydrogenase (EC 1.1.1.82) or fructose-1,6-bisphosphate phosphatase (EC 3.1.3.11) was assayed in a reconstituted chloroplastic, system comprising the isolated proteins of the ferredoxin-thioredoxin light-activation system and thylakoids from either mesophyll or bundle-sheath tissues of different C4 plants. While C4-plant thylakoids functionned almost equally well with C3-or C4-plant proteins, the photosyntem-II-deficient bundle-sheath thylakoids from the NADP-malic enzyme type, were unable to perform enzyme photoactivation unless supplemented with an electron donor to photosystem I. Bundle-sheath thylakoids isolated from plants showing no photosystem-II deficiency did not require such an addition. The results are discussed with respect to a possible requirement for a physiological reductant of ferredoxin for enzyme light activation in bundle-sheath, tissues.

Logarithmic phase Escherichia coli K1 efficiently avoids serum killing by promoting C4bp-mediated C3b and C4b degradation

PubMed Central

Wooster, David G; Maruvada, Ravi; Blom, Anna M; Prasadarao, Nemani V

2006-01-01

Meningitis caused by Escherichia coli K1 is a serious illness in neonates with neurological sequelae in up to 50% of survivors. A high degree of bacteremia is required for E. coli K1 to cross the blood–brain barrier, which suggests that the bacterium must evade the host defence mechanisms and survive in the bloodstream. We previously showed that outer membrane protein A (OmpA) of E. coli binds C4b-binding protein (C4bp), an inhibitor of complement activation via the classical pathway. Nevertheless, the exact mechanism by which E. coli K1 survives in serum remains elusive. Here, we demonstrate that log phase (LP) OmpA+E. coli K1 avoids serum bactericidal activity more effectively than postexponential phase bacteria. OmpA–E. coli cannot survive in serum grown to either phase. The increased serum resistance of LP OmpA+E. coli is the result of increased binding of C4bp, with a concomitant decrease in the deposition of C3b and the downstream complement proteins responsible for the formation of the membrane attack complex. C4bp bound to E. coli K1 acts as a cofactor to factor I in the cleavage of both C3b and C4b, which shuts down the ensuing complement cascade. Accordingly, a peptide corresponding to the complement control protein domain 3 of C4bp sequence, was able to compete with C4bp binding to OmpA and cause increased deposition of C3b. Thus, binding of C4bp appears to be responsible for survival of E. coli K1 in human serum. PMID:16556262

Isolated oxygen defects in 3C- and 4H-SiC: A theoretical study

NASA Astrophysics Data System (ADS)

Gali, A.; Heringer, D.; Deák, P.; Hajnal, Z.; Frauenheim, Th.; Devaty, R. P.; Choyke, W. J.

2002-09-01

Ab initio calculations in the local-density approximation have been carried out in SiC to determine the possible configurations of the isolated oxygen impurity. Equilibrium geometry and occupation levels were calculated. Substitutional oxygen in 3C-SiC is a relatively shallow effective mass like double donor on the carbon site (OC) and a hyperdeep double donor on the Si site (OSi). In 4H-SiC OC is still a double donor but with a more localized electron state. In 3C-SiC OC is substantially more stable under any condition than OSi or interstitial oxygen (Oi). In 4H-SiC OC is also the most stable one except for heavy n-type doping. We propose that OC is at the core of the electrically active oxygen-related defect family found by deep level transient spectroscopy in 4H-SiC. The consequences of the site preference of oxygen on the SiC/SiO2 interface are discussed.

C-Glycosyltransferases catalyzing the formation of di-C-glucosyl flavonoids in citrus plants.

PubMed

Ito, Takamitsu; Fujimoto, Shunsuke; Suito, Fumiaki; Shimosaka, Makoto; Taguchi, Goro

2017-07-01

Citrus plants accumulate many kinds of flavonoids, including di-C-glucosyl flavonoids, which have attracted considerable attention due to their health benefits. However, the biosynthesis of di-C-glucosyl flavonoids has not been elucidated at the molecular level. Here, we identified the C-glycosyltransferases (CGTs) FcCGT (UGT708G1) and CuCGT (UGT708G2) as the primary enzymes involved in the biosynthesis of di-C-glucosyl flavonoids in the citrus plants kumquat (Fortunella crassifolia) and satsuma mandarin (Citrus unshiu), respectively. The amino acid sequences of these CGTs were 98% identical, indicating that CGT genes are highly conserved in the citrus family. The recombinant enzymes FcCGT and CuCGT utilized 2-hydroxyflavanones, dihydrochalcone, and their mono-C-glucosides as sugar acceptors and produced corresponding di-C-glucosides. The K m and k cat values of FcCGT toward phloretin were <0.5 μm and 12.0 sec -1 , and those toward nothofagin (3'-C-glucosylphloretin) were 14.4 μm and 5.3 sec -1 , respectively; these values are comparable with those of other glycosyltransferases reported to date. Transcripts of both CGT genes were found to concentrate in various plant organs, and particularly in leaves. Our results suggest that di-C-glucosyl flavonoid biosynthesis proceeds via a single enzyme using either 2-hydroxyflavanones or phloretin as a substrate in citrus plants. In addition, Escherichia coli cells expressing CGT genes were found to be capable of producing di-C-glucosyl flavonoids, which is promising for commercial production of these valuable compounds. © 2017 The Authors The Plant Journal © 2017 John Wiley & Sons Ltd.

Elevation-induced climate change as a dominant factor causing the late Miocene C(4) plant expansion in the Himalayan foreland.

PubMed

Wu, Haibin; Guo, Zhengtang; Guiot, Joël; Hatté, Christine; Peng, Changhui; Yu, Yanyan; Ge, Junyi; Li, Qin; Sun, Aizhi; Zhao, Deai

2014-05-01

During the late Miocene, a dramatic global expansion of C4 plant distribution occurred with broad spatial and temporal variations. Although the event is well documented, whether subsequent expansions were caused by a decreased atmospheric CO2 concentration or climate change is a contentious issue. In this study, we used an improved inverse vegetation modeling approach that accounts for the physiological responses of C3 and C4 plants to quantitatively reconstruct the paleoclimate in the Siwalik of Nepal based on pollen and carbon isotope data. We also studied the sensitivity of the C3 and C4 plants to changes in the climate and the atmospheric CO2 concentration. We suggest that the expansion of the C4 plant distribution during the late Miocene may have been primarily triggered by regional aridification and temperature increases. The expansion was unlikely caused by reduced CO2 levels alone. Our findings suggest that this abrupt ecological shift mainly resulted from climate changes related to the decreased elevation of the Himalayan foreland. © 2013 John Wiley & Sons Ltd.

Finding the genes to build C4 rice.

PubMed

Wang, Peng; Vlad, Daniela; Langdale, Jane A

2016-06-01

Rice, a C3 crop, is a staple food for more than half of the world's population, with most consumers living in developing countries. Engineering C4 photosynthetic traits into rice is increasingly suggested as a way to meet the 50% yield increase that is predicted to be needed by 2050. Advances in genome-wide deep-sequencing, gene discovery and genome editing platforms have brought the possibility of engineering a C3 to C4 conversion closer than ever before. Because C4 plants have evolved independently multiple times from C3 origins, it is probably that key genes and gene regulatory networks that regulate C4 were recruited from C3 ancestors. In the past five years there have been over 20 comparative transcriptomic studies published that aimed to identify these recruited C4 genes and regulatory mechanisms. Here we present an overview of what we have learned so far and preview the efforts still needed to provide a practical blueprint for building C4 rice. Copyright © 2016 Elsevier Ltd. All rights reserved.

Middle to Late Holocene Fluctuations of C3 and C4 Vegetation in a Northern New England Salt Marsh, Sprague Marsh, Phippsburg Maine

DOE Office of Scientific and Technical Information (OSTI.GOV)

Johnson, B J; Moore, K A; Lehmann, C

2006-05-26

A 3.1 meter sediment core was analyzed for stable carbon isotope composition of organic matter and higher plant leaf wax (HPLW) lipid biomarkers to determine Holocene shifts in C{sub 3} (higher high marsh) and C{sub 4} (low and/or high marsh) plant deposition at the Sprague River Salt Marsh, Phippsburg, Maine. The carbon isotope composition of the bulk sediment and the HPLW parallel each other throughout most of the core, suggesting that terrestrial plants are an important source of organic matter to the sediments, and diagenetic alteration of the bulk sediments is minimal. The current salt marsh began to form 2500more » cal yr BP. Low and/or high C{sub 4} marsh plants dominated deposition at 2000 cal yr BP, 700 cal yr BP, and for the last 200 cal yr BP. Expansion of higher high marsh C{sub 3} plants occurred at 1300 and 600 cal yr BP. These major vegetation shifts result from a combination of changes in relative sea-level rise and sediment accumulation rates. Average annual carbon sequestration rates for the last 2500 years approximate 40 g C yr{sup -1} m{sup -2}, and are in strong agreement with other values published for the Gulf of Maine. Given that Maine salt marshes cover an area of {approx}79 km{sup 2}, they represent an important component of the terrestrial carbon sink. More detailed isotopic and age records from a network of sediment cores at Sprague Marsh are needed to truly evaluate the long term changes in salt marsh plant communities and the impact of more recent human activity, including global warming, on salt marsh vegetation.« less

Epitaxy of boron phosphide on AlN, 4H-SiC, 3C-SiC and ZrB2 substrates

NASA Astrophysics Data System (ADS)

Padavala, Balabalaji

The semiconductor boron phosphide (BP) has many outstanding features making it attractive for developing various electronic devices, including neutron detectors. In order to improve the efficiency of these devices, BP must have high crystal quality along with the best possible electrical properties. This research is focused on growing high quality crystalline BP films on a variety of superior substrates like AlN, 4H-SiC, 3C-SiC and ZrB2 by chemical vapor deposition. In particular, the influence of various parameters such as temperature, reactant flow rates, and substrate type and its crystalline orientation on the properties of BP films were studied in detail. Twin-free BP films were produced by depositing on off-axis 4H-SiC(0001) substrate tilted 4° toward [11¯00] and crystal symmetry matched zincblende 3C-SiC. BP crystalline quality improved at higher deposition temperature (1200°C) when deposited on AlN, 4H-SiC, whereas increased strain in 3C-SiC and increased boron segregation in ZrB2 at higher temperatures limited the best deposition temperature to below 1200°C. In addition, higher flow ratios of PH 3 to B2H6 resulted in smoother films and improved quality of BP on all substrates. The FWHM of the Raman peak (6.1 cm -1), XRD BP(111) peak FWHM (0.18°) and peak ratios of BP(111)/(200) = 5157 and BP(111)/(220) = 7226 measured on AlN/sapphire were the best values reported in the literature for BP epitaxial films. The undoped films on AlN/sapphire were n-type with a highest electron mobility of 37.8 cm2/V˙s and a lowest carrier concentration of 3.15x1018 cm -3. Raman imaging had lower values of FWHM (4.8 cm-1 ) and a standard deviation (0.56 cm-1) for BP films on AlN/sapphire compared to 4H-SiC, 3C-SiC substrates. X-ray diffraction and Raman spectroscopy revealed residual tensile strain in BP on 4H-SiC, 3C-SiC, ZrB2/4H-SiC, bulk AlN substrates while compressive strain was evident on AlN/sapphire and bulk ZrB2 substrates. Among the substrates studied, Al

Synthesis and characterization of 3-ketohexadecanoic acid-1-14-C, DL-3-hydroxyhexadecanoic acid-1-14-C, and trans-2-hexadecenoic acid-1-14-C.

PubMed

Jones, J A; Blecher, M

1966-05-01

The chemical synthesis and characterization of three intermediates in the Beta oxidation of palmitic acid-1-(14)C by rat liver mitochondria, namely, 3-ketohexadecanoic acid-1-(14)C, DL-3-hydroxyhexadecanoic acid-1-(14)C, and trans-2-hexadecenoic acid-1-(14)C, are described.

Genome-scale modeling of the evolutionary path to C4 photosynthesis

NASA Astrophysics Data System (ADS)

Myers, Christopher R.; Bogart, Eli

In C4 photosynthesis, plants maintain a high carbon dioxide level in specialized bundle sheath cells surrounding leaf veins and restrict CO2 assimilation to those cells, favoring CO2 over O2 in competition for Rubisco active sites. In C3 plants, which do not possess such a carbon concentrating mechanism, CO2 fixation is reduced due to this competition. Despite the complexity of the C4 system, it has evolved convergently from more than 60 independent origins in diverse families of plants around the world over the last 30 million years. We study the evolution of the C4 system in a genome-scale model of plant metabolism that describes interacting mesophyll and bundle sheath cells and enforces key nonlinear kinetic relationships. Adapting the zero-temperature string method for simulating transition paths in physics and chemistry, we find the highest-fitness paths connecting C3 and C4 positions in the model's high-dimensional parameter space, and show that they reproduce known aspects of the C3-C4 transition while making additional predictions about metabolic changes along the path. We explore the relationship between evolutionary history and C4 biochemical subtype, and the effects of atmospheric carbon dioxide levels.

Evidence of shift in C4 species range in central Argentina during the late Holocene

USGS Publications Warehouse

Silva, L.C.R.; Giorgis, M.A.; Anand, M.; Enrico, L.; Perez-Harguindeguy, N.; Falczuk, V.; Tieszen, L.L.; Cabido, M.

2011-01-01

AimMillennial-scale biogeographic changes are well understood in many parts of the world, but little is known about long-term vegetation dynamics in subtropical regions. Here we investigate shifts in C3/C4 plant abundance occurred in central Argentina during the past few millenniaMethodsWe determined present day soil organic matter δ13C signatures of grasslands, shrublands and woodlands, containing different mixtures of C3 and C4 plants. We measured past changes in the relative cover of C3/C4 plants by comparing δ13C values in soil profiles with present day δ13C signatures. We analyzed 14C activity in soil depths that showed major changes in vegetation.ResultsPresent day relative cover of C3/C4 plants determines whole ecosystem δ13C signatures integrated as litter and superficial soil organic matter (R2 = 0.78; p < 0.01). Deeper soils show a consistent shift in δ13C, indicating a continuous replacement of C4 by C3 plants since 3,870 (±210) YBP. During this period, the relative abundance of C3 plants increased 32% (average across sites) with significant changes being observed in all studied ecosystems.ConclusionsOur results show that C4 species were more abundant in the past, but C3 species became dominant during the late Holocene. We identified increases in the relative C3/C4 cover in grasslands, shrublands and woodlands, suggesting a physiological basis for changes in vegetation. The replacement of C4 by C3 plants coincided with changes in climate towards colder and wetter conditions and could represent a climatically driven shift in the C4 species optimum range.

Late-Quaternary variation in C3 and C4 grass abundance in southeastern Australia as inferred from δ13C analysis: Assessing the roles of climate, pCO2, and fire

NASA Astrophysics Data System (ADS)

Nelson, David M.; Urban, Michael A.; Kershaw, A. Peter; Hu, Feng Sheng

2016-05-01

Climate, atmospheric pCO2, and fire all may exert major influences on the relative abundance of C3 and C4 grasses in the present-day vegetation. However, the relative role of these factors in driving variation in C3 and C4 grass abundances in the paleorecord is uncertain, and C4 abundance is often interpreted narrowly as a proxy indicator of aridity or pCO2. We measured δ13C values of individual grains of grass (Poaceae) pollen in the sediments of two sites in southeastern Australia to assess changes in the proportions of C3 and C4 grasses during the past 25,000 years. These data were compared with shifts in pCO2, temperature, moisture balance, and fire to assess how these factors were related to long-term variation of C4 grass abundance during the late Quaternary. At Caledonia Fen, a high-elevation site in the Snowy Mountains, C4 grass abundance decreased from an average of 66% during the glacial period to 11% during the Holocene, primarily in response to increased pCO2 and temperature. In contrast, this pattern did not exist in low-elevation savannah woodlands around Tower Hill Northwest Crater, where C4 grass abundance instead varied in response to shifts in regional aridity. Fire did not appear to have strongly influenced the proportions of C3 and C4 grasses on the landscape at millennial timescales at either site. These patterns are similar to those of a recent study in East Africa, suggesting that elevation-related climatic differences influence how the abundance of C3 and C4 grasses responds to shifts in climate and pCO2. These results caution against using C4 plant abundance as a proxy indicator of either climate or pCO2 without an adequate understanding of key controlling factors.

Two-Dimensional Nb-Based M 4 C 3 Solid Solutions (MXenes)

DOE PAGES

Yang, Jian; Naguib, Michael; Ghidiu, Michael; ...

2015-10-15

Two new two-dimensional Nb 4C 3-based solid solutions (MXenes), (Nb 0.8,Ti 0.2) 4C 3T x and (Nb 0.8,Zr 0.2) 4C 3T x (where T is a surface termination) were synthesizedas confirmed by X-ray diffractionfrom their corresponding MAX phase precursors (Nb 0.8,Ti 0.2) 4AlC 3 and (Nb 0.8,Zr 0.2) 4AlC 3. In our report we discuss Zr-containing MXene. We also studied intercalation of Li ions into these two compositions, and Nb 4C 3T x in order to determine the potential of those materials for energy storage applications. Lithiation and delithiation peaks at 2.26 and 2.35 V, respectively, appeared in the casemore » of Nb 4C 3T x, but were not present in Nb 2CT x. After 20 cycles at a rate of C/4, the specific capacities of (Nb 0.8,Ti 0.2) 4C 3T xand (Nb 0.8,Ti 0.2) 4C 3T x were 158 and 132 mAh/g, respectively, both slightly lower than the capacity of Nb 4C 3T x.« less

Arabidopsis C3HC4-RING finger E3 ubiquitin ligase AtAIRP4 positively regulates stress-responsive abscisic acid signaling.

PubMed

Yang, Liang; Liu, Qiaohong; Liu, Zhibin; Yang, Hao; Wang, Jianmei; Li, Xufeng; Yang, Yi

2016-01-01

Degradation of proteins via the ubiquitin system is an important step in many stress signaling pathways in plants. E3 ligases recognize ligand proteins and dictate the high specificity of protein degradation, and thus, play a pivotal role in ubiquitination. Here, we identified a gene, named Arabidopsis thaliana abscisic acid (ABA)-insensitive RING protein 4 (AtAIRP4), which is induced by ABA and other stress treatments. AtAIRP4 encodes a cellular protein with a C3HC4-RING finger domain in its C-terminal side, which has in vitro E3 ligase activity. Loss of AtAIRP4 leads to a decrease in sensitivity of root elongation and stomatal closure to ABA, whereas overexpression of this gene in the T-DNA insertion mutant atairp4 effectively recovered the ABA-associated phenotypes. AtAIRP4 overexpression plants were hypersensitive to salt and osmotic stresses during seed germination, and showed drought avoidance compared with the wild-type and atairp4 mutant plants. In addition, the expression levels of ABA- and drought-induced marker genes in AtAIRP4 overexpression plants were markedly higher than those in the wild-type and atairp4 mutant plants. Hence, these results indicate that AtAIRP4 may act as a positive regulator of ABA-mediated drought avoidance and a negative regulator of salt tolerance in Arabidopsis. © 2015 The Authors. Journal of Integrative Plant Biology published by Wiley Publishing Asia Pty Ltd on behalf of Institute of Botany, Chinese Academy of Sciences.

Elementary reaction profile and chemical kinetics study of [C(1D)/(3P) + SiH4] with the CCSD(T) method

NASA Astrophysics Data System (ADS)

Ranka, Karnamohit; Perera, Ajith; Bartlett, Rodney J.

2017-07-01

Carbon and silicon-based molecules are omnipresent in the fields of combustion, atmospheric, semiconductor, and astronomical chemistry, among others. This paper reports the underlying elementary reactions for the [C(1D) + SiH4] and [C(3P) + SiH4] reaction profiles, optimized geometries of the intermediates, transition states (at the CCSD(T) level), RRKM and TST rate constants, and the corresponding branching ratios. Previously unreported van der Waals complex intermediates have been found for both reactions.

Effects of different elevated CO2 concentrations on chlorophyll contents, gas exchange, water use efficiency, and PSII activity on C3 and C4 cereal crops in a closed artificial ecosystem.

PubMed

Wang, Minjuan; Xie, Beizhen; Fu, Yuming; Dong, Chen; Hui, Liu; Guanghui, Liu; Liu, Hong

2015-12-01

Although terrestrial CO2 concentrations [CO2] are not expected to reach 1000 μmol mol(-1) (or ppm) for many decades, CO2 levels in closed systems such as growth chambers and greenhouses can easily exceed this concentration. CO2 levels in life support systems (LSS) in space can exceed 10,000 ppm (1 %). In order to understand how photosynthesis in C4 plants may respond to elevated CO2, it is necessary to determine if leaves of closed artificial ecosystem grown plants have a fully developed C4 photosynthetic apparatus, and whether or not photosynthesis in these leaves is more responsive to elevated [CO2] than leaves of C3 plants. To address this issue, we evaluated the response of gas exchange, water use efficiency, and photosynthetic efficiency of PSII by soybean (Glycine max (L.) Merr., 'Heihe35') of a typical C3 plant and maize (Zea mays L., 'Susheng') of C4 plant under four CO2 concentrations (500, 1000, 3000, and 5000 ppm), which were grown under controlled environmental conditions of Lunar Palace 1. The results showed that photosynthetic pigment by the C3 plants of soybean was more sensitive to elevated [CO2] below 3000 ppm than the C4 plants of maize. Elevated [CO2] to 1000 ppm induced a higher initial photosynthetic rate, while super-elevated [CO2] appeared to negate such initial growth promotion for C3 plants. The C4 plant had the highest ETR, φPSII, and qP under 500-3000 ppm [CO2], but then decreased substantially at 5000 ppm [CO2] for both species. Therefore, photosynthetic down-regulation and a decrease in photosynthetic electron transport occurred by both species in response to super-elevated [CO2] at 3000 and 5000 ppm. Accordingly, plants can be selected for and adapt to the efficient use of elevated CO2 concentration in LSS.

C4 plant isotopic composition (delta13C) evidence for urban CO2 pollution in the city of Cotonou, Benin (West Africa).

PubMed

Kèlomé, Nelly C; Lévêque, Jean; Andreux, Francis; Milloux, Marie-Jeanne; Oyédé, Lucien-Marc

2006-08-01

The carbon isotopic composition (delta13C) of plants can reveal the isotopic carbon content of the atmosphere in which they develop. The delta13C values of air and plants depend on the amount of atmospheric fossil fuel CO2, which is chiefly emitted in urban areas. A new indicator of CO2 pollution is tested using the delta13C variation in a C4 grass: Eleusine indica. A range of about 4 per thousand delta units was observed at different sites in Cotonou, the largest city in the Republic of Benin. The highest delta13C values, from -12 per thousand to -14 per thousand, were found in low traffic zones; low delta13C values, from -14 per thousand to -16 per thousand, were found in high traffic zones. The amount of fossil fuel carbon assimilated by plants represented about 20% of the total plant carbon content. An overall decrease in plant delta13C values was observed over a four-year monitoring period. This decrease was correlated with increasing vehicle traffic. The delta13C dataset and the corresponding geographical database were used to map and define zones of high and low 13C-depleted CO2 emissions in urban and sub-urban areas. The spatial distribution follows dominant wind directions, with the lowest emission zones found in the southwest of Cotonou. High CO2 emissions occurred in the north, the east and the center, providing evidence of intense anthropogenic activity related to industry and transportation.

Growth of Defect-Free 3C-SiC on 4H- and 6H-SiC Mesas Using Step-Free Surface Heteroepitaxy

NASA Technical Reports Server (NTRS)

Neudeck, Philip G.; Powell, J. Anthony; Trunek, Andrew J.; Huang, Xianrong R.; Dudley, Michael

2001-01-01

A new growth process, herein named step-free surface heteroepitaxy, has achieved 3CSiC films completely free of double positioning boundaries and stacking faults on 4H-SiC and 6H-SiC substrate mesas. The process is based upon the initial 2-dimensional nucleation and lateral expansion of a single island of 3C-SiC on a 4H- or 6H-SiC mesa surface that is completely free of bilayer surface steps. Our experimental results indicate that substrate-epilayer in-plane lattice mismatch (delta a/a = 0.0854% for 3C/4H) is at least partially relieved parallel to the interface in the initial bilayers of the heterofilm, producing an at least partially relaxed 3C-SiC film without dislocations that undesirably thread through the thickness of the epilayer. This result should enable realization of improved 3C-SiC devices.

Atmospheric Oxygen Concentrations for the Past 350 Myr Modeled from the δ13C of C3 Land Plants

NASA Astrophysics Data System (ADS)

Nordt, L.; Breecker, D.

2016-12-01

Numerous studies have focused on the systematic collection of long-term d13C records from marine sediments, but no such isotopic compilation exists for C3 land plants. Consequently, we gathered a meta-data base of 8003 plant-derived δ13C values (ISOORG) from various carbon sources binned into 5 myr time steps. The results of this investigation were reported in a recent publication showing that most δ13C sources co-vary with ten CIEs during the last 400 myr. For this paper we culled ISOORG to produce ISOORG16-H that contains 7025 plant-derived δ13C values from paleo-moist environments to reflect secular controls on the δ13C of C3 plants. We then constructed atmospheric pO2 curves for the past 350 myr using prior experimental work showing a direct relationship between the Δ13C of C3 plants and pO2 concentration. Periods of hyperoxia (25-30% pO2) were identified from 300-250, 225-190, and 110-105 myr, and intervals of hypoxia (10-15% pO2) from 350-345, 245-230, and 185-115 myr. During the last 150 myr, pO2 stabilized at 17-24% except for a notable positive excursion from 110-105 myr. Hyperoxia, apparently from widespread carbon burial, supports the notion of insect gigantism during the Late Paleozoic. Hypoxia during the early Triassic correlates with the coal gap following the collapse of Paleozoic ecosystems. Rising pO2 in the late Triassic seems to reflect renewed carbon burial from reorganization of Mesozoic ecosystems. The middle Mesozoic is characterized by low pO2 during an intense greenhouse interval, with ambient conditions ensuing thereafter possibly linked to carbon burial from the radiation of angiosperms. pO2 concentrations >14% suggest wildfires persisted through the study interval except possibly at 160 and 140 myr. Intervals of low pO2 concentration were likely accompanied by lower atmospheric pressure and higher temperatures, particularly from 245-230 myr and 180-120 myr. Our O2 reconstructions conform with GEOCARBSULF, but not with proxy

X-Ray study of hetero ring flexibility in norbornane, norbornene-fused 1,3-oxazin-2-thiones Structure of 5,8-methano- r-4-phenyl-c-4a, c-5,6,7, c-8, c- 8a-hexahydro-4H- 1 ,3-benzoxazin-2 (3H)-thione

NASA Astrophysics Data System (ADS)

Kálmán, A.; Argay, Gy.; Stájer, G.; Bernáth, G.

1991-08-01

The structure of S,8-methano- r-4-phenyl c4a, c5,6,7 c8 c8ahexahydro4 H 1,3 -benzoxazin- 2 (3 H)-thione (C 15H 17N0S, M r=259.37) has been established by X-ray crystallography from diffractometer data: it crystallizes in the monoclinic space group P2 1/n with a=6.150(2) Å, b=9.655(1) Å, c=22.093(4) Å,β=96.75(2)† V=1302.7(8) Å 3,4,D c=1.32gcm -3and p( Cu K) =20.4cm -. The structure has been solved by direct methods, refined to R=0.050 for 2193 observed reflections. The X-ray analysis substantiated the structure: the NMR spectra in- dicated that the 4-phenyl group assumes an exo-equatorial position. The puckering parameters of D. Cremer, J.A. Pople, J. Am. Chem. Soc., 97 (1975), 1354 (ref.1), of the distorted hetero ring (a transitional form between E 4-envelope, ( 5S 4-screw-boat) show that, depending on the positions of the hetero atoms, both the norbornane, norbornene skeletons markedly alter the characteristic transitional ( 5E/ 5H 6) shape of the 1,3-oxazine ring observed in other saturated, partly saturated l,3-benzoxazin-2-ones, analogous thiones.

C/sub 4/ photosynthesis in Euphorbia degeneri and E. remyi: a comparison of photosynthetic carbon metabolism in leaves, callus cultures and regenerated plants

DOE Office of Scientific and Technical Information (OSTI.GOV)

Ruzin, S.E.

1984-04-01

Based on analysis of /sup 14/CO/sub 2/ fixation kinetics and assays of enzymes related to C/sub 4/ metabolism (NAD-ME, NADP-ME, NAD-MDH, NADP-MDH, AST, ALT), leaves and regenerated plants of Euphorbia degeneri exhibit a modified NADP-ME-type photosynthesis. Apparently, both aspartate and malate are used for transport of CO/sub 2/ to bundle sheath cells. Callus grown on either non-shoot-forming or shoot-forming media fixes CO/sub 2/ into RPP-cycle intermediates and sucrose, as well as malate and aspartate. /sup 14/CO/sub 2/ pulse/chase kinetics show no significant loss of label from C/sub 4/ acids throughout a one minute chase. Analysis of PEPCase revealed the presencemore » of 2 isoenzymes in both leaf and regenerated plant tissues (K/sub m/ (PEP) = 0.080 and 0.550) but only one isoenzyme in callus (K/sub m/ = 0.100). It appears that C/sub 4/ photosynthesis does not occur in callus derived from this C/sub 4/ dicot but is regenerated concomitant with shoot regeneration, and ..beta..-carboxylation of PEP in callus, mediated by the low K/sub m/ isoenzyme of PEPCase, produces C/sub 4/ acids that are not involved in the CO/sub 2/ shuttle mechanism characteristic of C/sub 4/ photosynthesis. 161 references, 19 figures, 12 tables.« less

The Crystal Structure of Cobra Venom Factor, a Cofactor for C3- and C5-Convertase CVFBb

DOE Office of Scientific and Technical Information (OSTI.GOV)

Krishnan, Vengadesan; Ponnuraj, Karthe; Xu, Yuanyuan

2009-05-26

Cobra venom factor (CVF) is a functional analog of human complement component C3b, the active fragment of C3. Similar to C3b, in human and mammalian serum, CVF binds factor B, which is then cleaved by factor D, giving rise to the CVFBb complex that targets the same scissile bond in C3 as the authentic complement convertases C4bC2a and C3bBb. Unlike the latter, CVFBb is a stable complex and an efficient C5 convertase. We solved the crystal structure of CVF, isolated from Naja naja kouthia venom, at 2.6 {angstrom} resolution. The CVF crystal structure, an intermediate between C3b and C3c, lacksmore » the TED domain and has the CUB domain in an identical position to that seen in C3b. The similarly positioned CUB and slightly displaced C345c domains of CVF could play a vital role in the formation of C3 convertases by providing important primary binding sites for factor B.« less

The crystal structure of cobra venom factor, a cofactor for C3- and C5-convertase CVFBb.

PubMed

Krishnan, Vengadesan; Ponnuraj, Karthe; Xu, Yuanyuan; Macon, Kevin; Volanakis, John E; Narayana, Sthanam V L

2009-04-15

Cobra venom factor (CVF) is a functional analog of human complement component C3b, the active fragment of C3. Similar to C3b, in human and mammalian serum, CVF binds factor B, which is then cleaved by factor D, giving rise to the CVFBb complex that targets the same scissile bond in C3 as the authentic complement convertases C4bC2a and C3bBb. Unlike the latter, CVFBb is a stable complex and an efficient C5 convertase. We solved the crystal structure of CVF, isolated from Naja naja kouthia venom, at 2.6 A resolution. The CVF crystal structure, an intermediate between C3b and C3c, lacks the TED domain and has the CUB domain in an identical position to that seen in C3b. The similarly positioned CUB and slightly displaced C345c domains of CVF could play a vital role in the formation of C3 convertases by providing important primary binding sites for factor B.

Evolution of CAM and C4 carbon-concentrating mechanisms

USGS Publications Warehouse

Keeley, Jon E.; Rundel, Philip W.

2003-01-01

Mechanisms for concentrating carbon around the Rubisco enzyme, which drives the carbon-reducing steps in photosynthesis, are widespread in plants; in vascular plants they are known as crassulacean acid metabolism (CAM) and C4 photosynthesis. CAM is common in desert succulents, tropical epiphytes, and aquatic plants and is characterized by nighttime fixation of CO2. The proximal selective factor driving the evolution of this CO2-concentrating pathway is low daytime CO2, which results from the unusual reverse stomatal behavior of terrestrial CAM species or from patterns of ambient CO2 availability for aquatic CAM species. In terrestrials the ultimate selective factor is water stress that has selected for increased water use efficiency. In aquatics the ultimate selective factor is diel fluctuations in CO2 availability for palustrine species and extreme oligotrophic conditions for lacustrine species. C4 photosynthesis is based on similar biochemistry but carboxylation steps are spatially separated in the leaf rather than temporally as in CAM. This biochemical pathway is most commonly associated with a specialized leaf anatomy known as Kranz anatomy; however, there are exceptions. The ultimate selective factor driving the evolution of this pathway is excessively high photorespiration that inhibits normal C3 photosynthesis under high light and high temperature in both terrestrial and aquatic habitats. CAM is an ancient pathway that likely has been present since the Paleozoic era in aquatic species from shallow-water palustrine habitats. While atmospheric CO2 levels have undoubtedly affected the evolution of terrestrial plant carbon-concentrating mechanisms, there is reason to believe that past atmospheric changes have not played as important a selective role in the aquatic milieu since palustrine habitats today are not generally carbon sinks, and the selective factors driving aquatic CAM are autogenic. Terrestrial CAM, in contrast, is of increasing selective value under

Registration of NE Trailblazer C-1, NE Trailblazer C0, NE Trailblazer C2, NE Trailblazer C3, NE Trailblazer C4, and NE Trailblazer C5 Switchgrass Germplasms

USDA-ARS?s Scientific Manuscript database

NE Trailblazer C-1 (GP-101, PI 672015), NE Trailblazer C0 (GP-100, PI 672014), NE Trailblazer C2 (GP-102, PI 672016), NE Trailblazer C3 (GP-103, PI 672017), NE Trailblazer C4 (GP-104, PI 672018), and NE Trailblazer C5 (GP-105, PI 672019) switchgrass (Panicum virgatum L.) germplasms were released by ...

Isolation, observation, and computational modeling of proposed intermediates in catalytic proton reductions with the hydrogenase mimic Fe2(CO)6S2C6H4.

PubMed

Wright, Robert J; Zhang, Wei; Yang, Xinzheng; Fasulo, Meg; Tilley, T Don

2012-01-07

Proposed electrocatalytic proton reduction intermediates of hydrogenase mimics were synthesized, observed, and studied computationally. A new mechanism for H(2) generation appears to involve Fe(2)(CO)(6)(1,2-S(2)C(6)H(4)) (3), the dianions {[1,2-S(2)C(6)H(4)][Fe(CO)(3)(μ-CO)Fe(CO)(2)](2-) (3(2-)), the bridging hydride {[1,2-S(2)C(6)H(4)][Fe(CO)(3)(μ-CO)(μ-H)Fe(CO)(2)]}(-), 3H(-)(bridging), and the terminal hydride 3H(-)(term-stag), {[1,2-S(2)C(6)H(4)][HFe(CO)(3)Fe(CO)(3)]}(-), as intermediates. The dimeric sodium derivative of 3(2-), {[Na(2)(THF)(OEt(2))(3)][3(2-)]}(2) (4) was isolated from reaction of Fe(2)(CO)(6)(1,2-S(2)C(6)H(4)) (3) with excess sodium and was characterized by X-ray crystallography. It possesses a bridging CO and an unsymmetrically bridging dithiolate ligand. Complex 4 reacts with 4 equiv. of triflic or benzoic acid (2 equiv. per Fe center) to generate H(2) and 3 in 75% and 60% yields, respectively. Reaction of 4 with 2 equiv. of benzoic acid generated two hydrides in a 1.7 : 1 ratio (by (1)H NMR spectroscopy). Chemical shift calculations on geometry optimized structures of possible hydride isomers strongly suggest that the main product, 3H(-)(bridging), possesses a bridging hydride ligand, while the minor product is a terminal hydride, 3H(-)(term-stag). Computational studies support a catalytic proton reduction mechanism involving a two-electron reduction of 3 that severs an Fe-S bond to generate a dangling thiolate and an electron rich Fe center. The latter iron center is the initial site of protonation, and this event is followed by protonation at the dangling thiolate to give the thiol thiolate [Fe(2)H(CO)(6)(1,2-SHSC(6)H(4))]. This species then undergoes an intramolecular acid-base reaction to form a dihydrogen complex that loses H(2) and regenerates 3.

Synthesis and characterization of a new and electronically unusual uranium metallacyclocumulene, (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph)

DOE PAGES

Pagano, Justin K.; Erickson, Karla A.; Scott, Brian L.; ...

2016-10-22

A new uranium metallacyclocumulene, (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph), was synthesized by both reaction of (C 5Me 5) 2UCl 2 with 1,4-diphenylbutadiyne in the presence of KC 8 and by ligand exchange between (C 5Me 5) 2U(η 2-Me 3SiC 2SiMe 3) and 1,4-diphenylbutadiyne. Lastly, full characterization of (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph) is reported, including the solid-state structure. (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph) displays an unusually detailed UV–visible spectrum, which is rare for uranium(IV) metallocene complexes.

Synthesis and characterization of a new and electronically unusual uranium metallacyclocumulene, (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph)

DOE Office of Scientific and Technical Information (OSTI.GOV)

Pagano, Justin K.; Erickson, Karla A.; Scott, Brian L.

A new uranium metallacyclocumulene, (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph), was synthesized by both reaction of (C 5Me 5) 2UCl 2 with 1,4-diphenylbutadiyne in the presence of KC 8 and by ligand exchange between (C 5Me 5) 2U(η 2-Me 3SiC 2SiMe 3) and 1,4-diphenylbutadiyne. Lastly, full characterization of (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph) is reported, including the solid-state structure. (C 5Me 5) 2U(η 4-1,2,3,4-PhC 4Ph) displays an unusually detailed UV–visible spectrum, which is rare for uranium(IV) metallocene complexes.

Field nanoemitter: One-dimension Al4C3 ceramics

NASA Astrophysics Data System (ADS)

Sun, Y.; Cui, H.; Gong, L.; Chen, Jian; Shen, P. K.; Wang, C. X.

2011-07-01

As a kind of ionic (or salt-like) carbide, Al4C3 hardly any active functions have been found except for structure material purposes. However, considering the unique characteristic features of its crystal structure, we think Al4C3 in fact might have huge potential for exhibiting active functionality on field-emission application. Herein, we report for the first time the catalyst-free synthesis and excellent field emission properties of Al4C3 one-dimension (1-D) nanostructures. The 1-D nanostructures acting as cold electron emitters display excellent field emission performance with the turn-on field as low as 1.4-2.0 V μm-1 and the threshold field down to 4.2-4.4 V μm-1. Such emitters are technologically useful, because they can be easily fabricated on large substrates, and the synthesis process is simple and broadly applicable. The findings conceptually provide new opportunities for the application of Al4C3 ceramic material in vacuum microelectronic devices.

Self-association and domain rearrangements between complement C3 and C3u provide insight into the activation mechanism of C3.

PubMed

Li, Keying; Gor, Jayesh; Perkins, Stephen J

2010-10-01

Component C3 is the central protein of the complement system. During complement activation, the thioester group in C3 is slowly hydrolysed to form C3u, then the presence of C3u enables the rapid conversion of C3 into functionally active C3b. C3u shows functional similarities to C3b. To clarify this mechanism, the self-association properties and solution structures of C3 and C3u were determined using analytical ultracentrifugation and X-ray scattering. Sedimentation coefficients identified two different dimerization events in both proteins. A fast dimerization was observed in 50 mM NaCl but not in 137 mM NaCl. Low amounts of a slow dimerization was observed for C3u and C3 in both buffers. The X-ray radius of gyration RG values were unchanged for both C3 and C3u in 137 mM NaCl, but depend on concentration in 50 mM NaCl. The C3 crystal structure gave good X-ray fits for C3 in 137 mM NaCl. By randomization of the TED (thioester-containing domain)/CUB (for complement protein subcomponents C1r/C1s, urchin embryonic growth factor and bone morphogenetic protein 1) domains in the C3b crystal structure, X-ray fits showed that the TED/CUB domains in C3u are extended and differ from the more compact arrangement of C3b. This TED/CUB conformation is intermediate between those of C3 and C3b. The greater exposure of the TED domain in C3u (which possesses the hydrolysed reactive thioester) accounts for the greater self-association of C3u in low-salt conditions. This conformational variability of the TED/CUB domains would facilitate their interactions with a broad range of antigenic surfaces. The second dimerization of C3 and C3u may correspond to a dimer observed in one of the crystal structures of C3b.

Biosynthesis of steroidal alkaloids in Solanaceae plants: involvement of an aldehyde intermediate during C-26 amination.

PubMed

Ohyama, Kiyoshi; Okawa, Akiko; Moriuchi, Yuka; Fujimoto, Yoshinori

2013-05-01

The C-26 amino group of steroidal alkaloids, such as tomatine, is introduced during an early step of their biosynthesis from cholesterol. In the present study, the mechanism of C-26 amination was reinvestigated by administering stable isotope labeled compounds, such as (26,26,26,27,27,27-(2)H6)cholesterol during biosynthesis of tomatine, solanine and solasonine. The chemical compositions of tomatine and solanine so obtained were analyzed by LC-MS after administering the d6-cholesterol to a tomato seedling and a potato shoot, respectively. The resulting spectra indicated that two deuterium atoms were eliminated from C-26 of cholesterol during biosynthesis. Furthermore, administration of (6-(13)C(2)H3)mevalonate in combination with lovastatin to an eggplant seedling, followed by GC-MS analysis of solasodine after TMS derivatization established that two deuterium atoms were eliminated from C-26 of cholesterol during solasonine biosynthesis. These findings are in contrast to an earlier observation that one hydrogen atom was lost from C-26 during tomatidine biosynthesis, and suggest that C-26 nitrogen atom addition involves an aldehyde intermediate. Thus, it is proposed that the C-26 amination reaction that occurs during steroidal alkaloid biosynthesis proceeds by way of a transamination mechanism. Copyright © 2013 Elsevier Ltd. All rights reserved.

Bimetallic-organic framework derived porous Co3O4/Fe3O4/C-loaded g-C3N4 nanocomposites as non-enzymic electrocatalysis oxidization toward ascorbic acid, dopamine acid, and uric acid

NASA Astrophysics Data System (ADS)

Hu, Bin; Liu, Yongkang; Wang, Zhuo-Wei; Song, Yingpan; Wang, Minghua; Zhang, Zhihong; Liu, Chun-Sen

2018-05-01

We report on the synthesis of Co- and Fe-based bimetallic nanocatalysts embedded in mesoporous carbon and g-C3N4 nanosheets (denoted as Co3O4/Fe3O4/mC@g-C3N4) for selectively simultaneous determination of ascorbic acid (AA), dopamine acid (DA), and uric acid (UA). These electrocatalysts consisting of bimetallic Co-Fe alloy nanoparticles encapsulated in N-doped carbon matrix were prepared via pyrolysis of Co/Fe-MOFs after grinding with high amounts of melamine. Chemical/crystal structures suggest high contents of mesoporous carbon in calcinated Co3O4/Fe3O4/mC nanocomposites, which exhibited enhanced electrocatalytic activity toward small biomolecules. The intrinsic performances of Co/Fe-MOFs with large specific surface area and regular nodes in the two-dimensional nanostructured g-C3N4 nanosheets endowed the as-prepared series of Co3O4/Fe3O4/mC@g-C3N4 nanocomposites with remarkable electrocatalytic activities and high adsorption ability toward oxidation of AA, DA, and UA. The developed biosensors also showed long-term stability and high selectivity for targeted analytes, with satisfactory results on actual samples in human urine. The results indicate that the as-synthesized Co3O4/Fe3O4/mC@g-C3N4 nanostructure exhibits good electrocatalytic activity and potential applications in clinical diagnosis and biosensing.

Late Pleistocene C4 plant dominance and summer rainfall in the southwestern United States from isotopic study of herbivore teeth

USGS Publications Warehouse

Connin, S.L.; Betancourt, J.; Quade, Jay

1998-01-01

Patterns of climate and C4 plant abundance in the southwestern United States during the last glaciation were evaluated from isotopic study of herbivore tooth enamel. Enamel ??13C values revealed a substantial eastward increase in C4 plant consumption for Mammuthus spp., Bison spp., Equus spp., and Camelops spp. The ??13C values were greatest in Bison spp. (-6.9 to + 1.7???) and Mammuthus spp. (-9.0 to +0.3???), and in some locales indicated C4-dominated grazing. The ??13C values of Antilocaprids were lowest among taxa (-12.5 to -7.9???) and indicated C3 feeding at all sites. On the basis of modern correlations between climate and C4 grass abundance, the enamel data imply significant summer rain in parts of southern Arizona and New Mexico throughout the last glaciation. Enamel ??18O values range from +19.0 to +31.0??? and generally increase to the east. This pattern could point to a tropical or subtropical source of summer rainfall. At a synoptic scale, the isotope data indicate that interactions of seasonal moisture, temperature, and lowered atmospheric pCO2 determined glacial-age C4 abundance patterns.

Effect of pH on the adsorption and photocatalytic degradation of sulfadimidine in Vis/g-C3N4 progress.

PubMed

Yang, Bin; Mao, Xuhui; Pi, Liu; Wu, Yixiao; Ding, Huijun; Zhang, Weihao

2017-03-01

In this study, g-C 3 N 4 was synthesized by thermal polycondensation of melamine and was characterized by X-ray powder diffraction, X-ray photoelectron spectroscopy, UV-visible diffuse reflection spectroscopy, and scanning electron microscopy. Results showed that g-C 3 N 4 degraded sulfadimidine (SMD) under visible light, in which the adsorption and photocatalytic degradation was influenced by pH. The maximum adsorption capacity was achieved at approximately pH 5. The highest degradation rate constant was obtained at strong acid and alkali. In addition, the degradation mechanism of g-C 3 N 4 was evaluated with the help of quencher agents. The intermediates, degradation pathways, and mineralization of SMD were also determined to evaluate the degradation and oxidation ability of g-C 3 N 4 .

Modeling C1-C4 Alkyl Nitrate Photochemistry and Their Impacts on O3 Production in Urban and Suburban Environments of Hong Kong

NASA Astrophysics Data System (ADS)

Lyu, X. P.; Guo, H.; Wang, N.; Simpson, I. J.; Cheng, H. R.; Zeng, L. W.; Saunders, S. M.; Lam, S. H. M.; Meinardi, S.; Blake, D. R.

2017-10-01

As intermediate products of photochemical reactions, alkyl nitrates (RONO2) regulate ozone (O3) formation. In this study, a photochemical box model incorporating master chemical mechanism well reproduced the observed RONO2 at an urban and a mountainous site, with index of agreement in the range of 0.66-0.73. The value 0.0003 was identified to be the most appropriate branching ratio for C1 RONO2, with the error less than 50%. Although levels of the parent hydrocarbons and nitric oxide (NO) were significantly higher at the urban site than the mountainous site, the production of C2-C3 RONO2 was comparable to or even lower than at the mountainous site, due to the lower concentrations of oxidative radicals in the urban environment. Based on the profiles of air pollutants at the mountainous site, the formation of C2-C4 RONO2 was limited by NOx (volatile organic compounds (VOCs)) when total volatile organic compounds (TVOCs)/NOx was higher (lower) than 10.0 ± 0.4 parts per billion by volume (ppbv)/ppbv. This dividing ratio decreased (p < 0.05) to 8.7 ± 0.4 ppbv/ppbv at the urban site, mainly due to the different air pollutant profiles at the two sites. For the formation of C1 RONO2, the NOx-limited regime extended the ratio of TVOCs/NOx to as low as 2.4 ± 0.2 and 3.1 ± 0.1 ppbv/ppbv at the mountainous and urban site, respectively. RONO2 formation led to a decrease of simulated O3, with reduction efficiencies (O3 reduction/RONO2 production) of 4-5 parts per trillion by volume (pptv)/pptv at the mountainous site and 3-4 pptv/pptv at the urban site. On the other hand, the variations of simulated O3 induced by RONO2 degradation depended upon the regimes controlling O3 formation and the relative abundances of TVOCs and NOx.

Comparative cell-specific transcriptomics reveals differentiation of C4 photosynthesis pathways in switchgrass and other C4 lineages

PubMed Central

Rao, Xiaolan; Lu, Nan; Li, Guifen; Nakashima, Jin; Tang, Yuhong; Dixon, Richard A.

2016-01-01

Almost all C4 plants require the co-ordination of the adjacent and fully differentiated cell types, mesophyll (M) and bundle sheath (BS). The C4 photosynthetic pathway operates through two distinct subtypes based on how malate is decarboxylated in BS cells; through NAD-malic enzyme (NAD-ME) or NADP-malic enzyme (NADP-ME). The diverse or unique cell-specific molecular features of M and BS cells from separate C4 subtypes of independent lineages remain to be determined. We here provide an M/BS cell type-specific transcriptome data set from the monocot NAD-ME subtype switchgrass (Panicum virgatum). A comparative transcriptomics approach was then applied to compare the M/BS mRNA profiles of switchgrass, monocot NADP-ME subtype C4 plants maize and Setaria viridis, and dicot NAD-ME subtype Cleome gynandra. We evaluated the convergence in the transcript abundance of core components in C4 photosynthesis and transcription factors to establish Kranz anatomy, as well as gene distribution of biological functions, in these four independent C4 lineages. We also estimated the divergence between NAD-ME and NADP-ME subtypes of C4 photosynthesis in the two cell types within C4 species, including differences in genes encoding decarboxylating enzymes, aminotransferases, and metabolite transporters, and differences in the cell-specific functional enrichment of RNA regulation and protein biogenesis/homeostasis. We suggest that C4 plants of independent lineages in both monocots and dicots underwent convergent evolution to establish C4 photosynthesis, while distinct C4 subtypes also underwent divergent processes for the optimization of M and BS cell co-ordination. The comprehensive data sets in our study provide a basis for further research on evolution of C4 species. PMID:26896851

A molten globule-like intermediate state detected in the thermal transition of cytochrome c under low salt concentration.

PubMed

Nakamura, Shigeyoshi; Baba, Takayuki; Kidokoro, Shun-Ichi

2007-04-01

To understand the stabilization mechanism of the transient intermediate state in protein folding, it is very important to understand the structure and stability of the molten globule state under a native condition, in which the native state exists stably. The thermal transitions of horse cytochrome c were thermodynamically evaluated by highly precise differential scanning calorimetry (DSC) at pH 3.8-5.0. The heat capacity functions were analyzed using double deconvolution and the nonlinear least-squares method. An intermediate (I) state is clearly confirmed in the thermal native (N)-to-denatured (D) transition of horse cytochrome c. The mole fraction of the intermediate state shows the largest value, 0.4, at nearly 70 degrees C at pH 4.1. This intermediate state was also detected by the circular dichroism (CD) method and was found to have the properties of the molten globule-like structure by three-state analysis of the CD data. The Gibbs free-energy change between N and I, DeltaG(NI), and that between N and D, DeltaG(ND), were evaluated to be 9-22 kJ mol(-1) and 41-45 kJ mol(-1), respectively at 15( ) degrees C and pH 4.1.

Two Novel C3N4 Phases: Structural, Mechanical and Electronic Properties

PubMed Central

Fan, Qingyang; Chai, Changchun; Wei, Qun; Yang, Yintang

2016-01-01

We systematically studied the physical properties of a novel superhard (t-C3N4) and a novel hard (m-C3N4) C3N4 allotrope. Detailed theoretical studies of the structural properties, elastic properties, density of states, and mechanical properties of these two C3N4 phases were carried out using first-principles calculations. The calculated elastic constants and the hardness revealed that t-C3N4 is ultra-incompressible and superhard, with a high bulk modulus of 375 GPa and a high hardness of 80 GPa. m-C3N4 and t-C3N4 both exhibit large anisotropy with respect to Poisson’s ratio, shear modulus, and Young’s modulus. Moreover, m-C3N4 is a quasi-direct-bandgap semiconductor, with a band gap of 4.522 eV, and t-C3N4 is also a quasi-direct-band-gap semiconductor, with a band gap of 4.210 eV, with the HSE06 functional. PMID:28773550

A Modified HR3C Austenitic Heat-Resistant Steel for Ultra-supercritical Power Plants Applications Beyond 650 °C

NASA Astrophysics Data System (ADS)

Zhu, C. Z.; Yuan, Y.; Zhang, P.; Yang, Z.; Zhou, Y. L.; Huang, J. Y.; Yin, H. F.; Dang, Y. Y.; Zhao, X. B.; Lu, J. T.; Yan, J. B.; You, C. Y.

2018-02-01

A modified HR3C austenitic steel has been designed by optimizing the chemical composition. Compared with a commercial HR3C alloy, the modified steel has comparable oxidation resistance, yield strength, and plasticity, but higher creep rupture strength and impact toughness after long-term thermal exposure. The results suggest that the modified alloy is a promising candidate for the applications of ultra-supercritical power plants operating beyond 650 °C.

Optional use of CAM photosynthesis in two C4 species, Portulaca cyclophylla and Portulaca digyna.

PubMed

Holtum, Joseph A M; Hancock, Lillian P; Edwards, Erika J; Winter, Klaus

2017-07-01

Low levels of crassulacean acid metabolism (CAM) are demonstrated in two species with C 4 photosynthesis, Portulaca cyclophylla and P. digyna. The expression of CAM in P. cyclophylla and P. digyna is facultative, i.e. optional. Well-watered plants did not accumulate acid at night and exhibited gas-exchange patterns consistent with C 4 photosynthesis. CAM-type nocturnal acidification was reversible in that it was induced following drought and lost when droughted plants were rewatered. In P. cyclophylla, droughting was accompanied by a small but discernible net uptake of CO 2 during the dark, whereas in P. digyna, net CO 2 exchange at night approached the CO 2 compensation point but did not transition beyond it. This report brings the number of known C 4 species with a capacity for expressing CAM to six. All are species of Portulaca. The observation of CAM in P. cyclophylla and P. digyna is the first for species in the opposite-leaved (OL) Portulacelloid-anatomy lineage of Portulaca and for the Australian clade therein. The other four species are within the alternate-leaved (AL) lineage, in the Atriploid-anatomy Oleracea and the Pilosoid-anatomy Pilosa clades. Studies of the evolutionary origins of C 4 and CAM in Portulaca will benefit from a more wide-range survey of CAM across its species, particularly in the C 3 -C 4 intermediate-containing Cryptopetala clade. Copyright © 2017 Elsevier GmbH. All rights reserved.

Mass spectral analysis of C3 and C4 aliphatic amino acid derivatives.

NASA Technical Reports Server (NTRS)

Lawless, J. G.; Chadha, M. S.

1971-01-01

Diagnostic criteria are obtained for the distinction of alpha, beta, gamma, and N-methyl isomers of the C3 and C4 aliphatic amino acids, using mass spectral analysis of the derivatives of these acids. The use of deuterium labeling has helped in the understanding of certain fragmentation pathways.

Comparison of physiological and anatomical changes of C3 (Oryza sativa [L.]) and C4 (Echinochloa crusgalli [L.]) leaves in response to drought stress

NASA Astrophysics Data System (ADS)

Hamim, Hamim; Banon, Sri; Dorly, Dorly

2016-01-01

The experiment aimed to analyse the different response of C3 (Oryza sativa L.) and C4 (Echinochloa crusgalli L.) species to drought stress based on physiological and anatomical properties. Seeds of rice (Oryza sativa) and Echinochloa (Echinochloa crusgalli) were grown in 15 cm (D) pot for 6 weeks under well-watered conditions. After 6 weeks the plants were divided into two groups, (1) well-watered which were watered daily, and (2) drought stress which were withheld from watering for 6 days. After 6 days of drought, the plants were then re-watered to analyse plant recovery. During drought period, the plants were analysed for growth, leaf relative water content (RWC), photosynthesis, and leaf anatomy. Drought stress significantly reduced leaf RWC of both species, but the reduction was bigger in rice than in Echinochloa. The maximum efficiency of photosynthesis (Fv/Fm) was decrease significantly in response to drought stress by about 48.04% in rice, while it was only 34.40% in Echinochloa. Anatomical analysis showed drought treatment tended to reduce leaf thickness in the area of bulliform cell, major- as well as intervein and xylem diameter, more in Echinochloa than in rice, suggesting that the decrease of vein and xylem diameter is among the anatomical parameters that is important to overcome from drought stress in Echinochloa. The number of chloroplast in the mesophyll cell and bundle sheath cell (BSC) was different between these two species, where in Echinochloa chloroplast was found in both mesophyll as well as BSC, while in rice it was only found in mesophyll cell, confirmed that Echinochloa is a C4 and rice is a C3 species. Interestingly, in Echinochloa, the number of chloroplast was significantly increased due to drought stress in BSC, but not in mesophyll cell. The number of starch granules also dramatically increased in response to drought in the mesophyll cells of rice and Echinochloa, and in the bundle sheath cell of Echinochloa which indicate that C3

Phase Equilibria and Transport Properties in the Systems AgNO3/RCN/H2O. R = CH3, C2H5, C3H7, C4H,, C6H5, and C6H5CH2

NASA Astrophysics Data System (ADS)

Das, Surjya P.; Wittekopf, Burghard; Weil, Konrad G.

1988-11-01

Silver nitrate can form homogeneous liquid phases with some organic nitriles and water, even when there is no miscibility between the pure liquid components. We determined the shapes of the single phase regions in the ternary phase diagram for the following systems: silver nitrate /RCN /H2O with R =CH3, C3H7, C6H5, and C6H5CH2 at room temperature and for R =C6H5 also at 60 °C and O °C. Furthermore we studied kinematic viscosities, electrical conductivities, and densities of mixtures containing silver nitrate, RCN, and water with the mole ratios X /4 /1 (0.2≦ X ≦S 3.4). In these cases also R = C2H5 and C4H9 were studied. The organic nitriles show different dependences of viscosity and conductivity on the silver nitrate content from the aliphatic ones.

Drought stress obliterates the preference for ammonium as an N source in the C4 plant Spartina alterniflora.

PubMed

Hessini, Kamel; Kronzucker, Herbert J; Abdelly, Chedly; Cruz, Cristina

2017-06-01

The C 4 grass Spartina alterniflora is known for its unique salt tolerance and strong preference for ammonium (NH 4 + ) as a nitrogen (N) source. We here examined whether Spartina's unique preference for NH 4 + results in improved performance under drought stress. Manipulative greenhouse experiments were carried out to measure the effects of variable water availability and inorganic N sources on plant performance (growth, photosynthesis, antioxidant, and N metabolism). Drought strongly reduced leaf number and area, plant fresh and dry weight, and photosynthetic activity on all N sources, but the reduction was most pronounced on NH 4 + . Indeed, the growth advantage seen on NH 4 + in the absence of drought, producing nearly double the biomass compared to growth on NO 3 - , was entirely obliterated under both intermediate and severe drought conditions (50 and 25% field capacity, respectively). Both fresh and dry weight became indistinguishable among N sources under drought. Major markers of the antioxidant capacity of the plant, the activities of the enzymes superoxide dismutase, catalase, ascorbate peroxidase, and glutathione reductase, showed higher constitutive levels on NH 4 + . Catalase and glutathione reductase were specifically upregulated in NH 4 + -fed plants with increasing drought stress. This upregulation, however, failed to protect the plants from drought stress. Nitrogen metabolism was characterized by lower constitutive levels of glutamine synthetase in NH 4 + -fed plants, and a rise in glutamate dehydrogenase (GDH) activity under drought, accompanied by elevated proline levels in leaves. Our results support postulates on the important role of GDH induction, and its involvement in the synthesis of compatible solutes, under abiotic stress. We show that, despite this metabolic shift, S. alterniflora's sensitivity to drought does not benefit from growth on NH 4 + and that the imposition of drought stress equalizes all N-source-related growth differences

Comparison of Ion Chemistries in Octafluoro-2-butene (2-C4F8) and in Octfluorocyclobutane (c-C4F8)

NASA Astrophysics Data System (ADS)

Jiao, Charles; Dejoseph, Charles; Garscadden, Alan

2007-10-01

2-C4F8 is one of the promising candidates to replace c-C4F8 that has been widely used for dielectric etching but is not environmentally friendly. In this study we have investigated electron impact ionization and ion-molecule reactions of 2-C4F8 using Fourier transform mass spectrometry (FTMS), and compared the results with those of c-C4F8 we have studied previously. Electron impact ionization of 2-C4F8 produces 15 ionic species including C4F7,8^+, C3F3,5,6^+, C2F4^+ and CF1-3^+ as the major ions. The total ionization cross section of 2-C4F8 reaches a maximum of 1.8x10-15 cm^2 at 90 eV. The ionization is dominated by the channel forming the parent ion C4F8^+ from 12 to 18 eV, and by the channel forming C3F5^+ from 18 to 70 eV. After 70 eV, CF3^+ becomes the dominant product ion. Among the major ions generated from the electron impact ionization of 2-C4F8, only CF^+, CF2^+ and CF3^+ are found to react with 2-C4F8, via F^- abstraction or charge transfer mechanism. The charge transfer reaction of Ar^++2-C4F8 produces primarily C4F7^+.

A novel RNA binding protein affects rbcL gene expression and is specific to bundle sheath chloroplasts in C4 plants

PubMed Central

2013-01-01

Background Plants that utilize the highly efficient C4 pathway of photosynthesis typically possess kranz-type leaf anatomy that consists of two morphologically and functionally distinct photosynthetic cell types, the bundle sheath (BS) and mesophyll (M) cells. These two cell types differentially express many genes that are required for C4 capability and function. In mature C4 leaves, the plastidic rbcL gene, encoding the large subunit of the primary CO2 fixation enzyme Rubisco, is expressed specifically within BS cells. Numerous studies have demonstrated that BS-specific rbcL gene expression is regulated predominantly at post-transcriptional levels, through the control of translation and mRNA stability. The identification of regulatory factors associated with C4 patterns of rbcL gene expression has been an elusive goal for many years. Results RLSB, encoded by the nuclear RLSB gene, is an S1-domain RNA binding protein purified from C4 chloroplasts based on its specific binding to plastid-encoded rbcL mRNA in vitro. Co-localized with LSU to chloroplasts, RLSB is highly conserved across many plant species. Most significantly, RLSB localizes specifically to leaf bundle sheath (BS) cells in C4 plants. Comparative analysis using maize (C4) and Arabidopsis (C3) reveals its tight association with rbcL gene expression in both plants. Reduced RLSB expression (through insertion mutation or RNA silencing, respectively) led to reductions in rbcL mRNA accumulation and LSU production. Additional developmental effects, such as virescent/yellow leaves, were likely associated with decreased photosynthetic function and disruption of associated signaling networks. Conclusions Reductions in RLSB expression, due to insertion mutation or gene silencing, are strictly correlated with reductions in rbcL gene expression in both maize and Arabidopsis. In both plants, accumulation of rbcL mRNA as well as synthesis of LSU protein were affected. These findings suggest that specific accumulation

Different growth responses of C3 and C4 grasses to seasonal water and nitrogen regimes and competition in a pot experiment.

PubMed

Niu, Shuli; Liu, Weixing; Wan, Shiqiang

2008-01-01

Understanding temporal niche separation between C(3) and C(4) species (e.g. C(3) species flourishing in a cool spring and autumn while C(4) species being more active in a hot summer) is essential for exploring the mechanism for their co-existence. Two parallel pot experiments were conducted, with one focusing on water and the other on nitrogen (N), to examine growth responses to water or nitrogen (N) seasonality and competition of two co-existing species Leymus chinensis (C(3) grass) and Chloris virgata (C(4) grass) in a grassland. The two species were planted in either monoculture (two individuals of one species per pot) or a mixture (two individuals including one L. chinensis and one C. virgata per pot) under three different water or N seasonality regimes, i.e. the average model (AM) with water or N evenly distributed over the growing season, the one-peak model (OPM) with more water or N in the summer than in the spring and autumn, and the two-peak model (TPM) with more water or N in the spring and autumn than in the summer. Seasonal water regimes significantly affected biomass in L. chinensis but not in C. virgata, while N seasonality impacted biomass and relative growth rate of both species over the growing season. L. chinensis accumulated more biomass under the AM and TPM than OPM water or N treatments. Final biomass of C. virgata was less impacted by water and N seasonality than that of L. chinensis. Interspecific competition significantly decreased final biomass in L. chinensis but not in C. virgata, suggesting an asymmetric competition between the two species. The magnitude of interspecific competition varied with water and N seasonality. Changes in productivity and competition balance of L. chinensis and C. virgata under shifting seasonal water and N availabilities suggest a contribution of seasonal variability in precipitation and N to the temporal niche separation between C(3) and C(4) species.

Carbon Isotope Discrimination in C3 Land Plants is Independent of Atmospheric PCO2

NASA Astrophysics Data System (ADS)

Kohn, M. J.

2015-12-01

The δ13C of terrestrial C3 plant tissues and soil organic matter is important for understanding the carbon cycle, inferring past climatic and ecological conditions, and predicting responses of vegetation to future climate change. Plant δ13C depends on the δ13C of atmospheric CO2 and mean annual precipitation (MAP), but an unresolved decades-long debate centers on whether terrestrial C3 plant δ13C responds to pCO2. Here, the pCO2-dependence of C3 land plant δ13C was tested using isotopic records from low- and high-pCO2 times spanning historical through Eocene data. Historical data do not resolve a clear pCO2-effect (-1.2±1.0 to 0.59±0.34‰/100 ppmv), and organic carbon records of the Pleistocene-Holocene transition implicate changes in MAP and ecosystems, rather than pCO2, as the major driver of δ13C changes. Fossil collagen and tooth enamel data constrain pCO2-effects most tightly to -0.03±0.13 and -0.03±0.24‰/100 ppmv between 200 and 700 ppmv. Combining all constraints yields a preferred value of 0.0±0.2‰/100 ppmv (2 s.e.), i.e. there is effectively no pCO2 effect. Recent models of pCO2-dependence imply unrealistic MAP for Cenozoic records.

Evolution of C4 plants: a new hypothesis for an interaction of CO2 and water relations mediated by plant hydraulics

PubMed Central

Osborne, Colin P.; Sack, Lawren

2012-01-01

C4 photosynthesis has evolved more than 60 times as a carbon-concentrating mechanism to augment the ancestral C3 photosynthetic pathway. The rate and the efficiency of photosynthesis are greater in the C4 than C3 type under atmospheric CO2 depletion, high light and temperature, suggesting these factors as important selective agents. This hypothesis is consistent with comparative analyses of grasses, which indicate repeated evolutionary transitions from shaded forest to open habitats. However, such environmental transitions also impact strongly on plant–water relations. We hypothesize that excessive demand for water transport associated with low CO2, high light and temperature would have selected for C4 photosynthesis not only to increase the efficiency and rate of photosynthesis, but also as a water-conserving mechanism. Our proposal is supported by evidence from the literature and physiological models. The C4 pathway allows high rates of photosynthesis at low stomatal conductance, even given low atmospheric CO2. The resultant decrease in transpiration protects the hydraulic system, allowing stomata to remain open and photosynthesis to be sustained for longer under drying atmospheric and soil conditions. The evolution of C4 photosynthesis therefore simultaneously improved plant carbon and water relations, conferring strong benefits as atmospheric CO2 declined and ecological demand for water rose. PMID:22232769

LC-MS/MS quantification of 7α-hydroxy-4-cholesten-3-one (C4) in rat and monkey plasma.

PubMed

Kang, Lijuan; Connolly, Thomas M; Weng, Naidong; Jian, Wenying

2017-10-01

7α-hydroxy-4-cholesten-3-one (C4) is an oxidative enzymatic product of cholesterol metabolism via cholesterol 7α-hydroxylase, an enzyme also known as cholesterol 7-alpha-monooxygenase or cytochrome P450 7A1 (CYP7A1). C4 is a stable intermediate in the rate limiting pathway of bile acid biosynthesis. Previous studies showed that plasma C4 levels correlated with CYP7A1 enzymatic activity and could serve as a biomarker for bile acid synthesis. Here we developed and qualified a simple and robust high-throughput method using liquid chromatography coupled with tandem mass spectrometry (LC-MS/MS) to quantify C4 in rat and monkey plasma. As C4 being an endogenous compound, this method used calibration standards in 50/50: acetonitrile/water (v/v). In order to mimic the incurred samples, quality control samples were prepared in the authentic plasma. Stable isotope labeled C4 (C4-d 7 ) was used as the internal standard. The sample volume for analysis was 20μL and the sample preparation method was protein precipitation with acetonitrile. The average endogenous C4 concentrations, from 10 different lots of rat and monkey plasma, were 53.0±16.5ng/mL and 6.8±5.6ng/mL, respectively. Based on these observed endogenous C4 levels, the calibration curve ranges were established at 1-200ng/mL and 0.5-100ng/mL for rat assay and monkey assay, respectively. The method was qualified with acceptable accuracy, precision, linearity, and specificity. Matrix effect, recovery, and plasma stability of bench-top, freeze-thaw, and long-term frozen storage were also evaluated. The method has been successfully applied to pre-clinical studies. Copyright © 2017 Elsevier B.V. All rights reserved.

Downregulation of p-coumaroyl quinate/shikimate 3'-hydroxylase (C3'H) or cinnamate-4-hydrolylase (C4H) in Eucalyptus urophylla x Eucalyptus grandis leads to increased extractability

DOE PAGES

Ziebell, Angela; Gjersing, Erica; Hinchee, Maud; ...

2016-01-20

Lignin reduction through breeding and genetic modification has the potential to reduce costs in biomass processing in pulp and paper, forage, and lignocellulosic ethanol industries. Here, we present detailed characterization of the extractability and lignin structure of Eucalyptus urophylla x Eucalyptus grandis RNAi downregulated in p-coumaroyl quinate/shikimate 3'-hydroxylase (C3'H) or cinnamate-4-hydroxylase (C4H). Both the C3'H and C4H downregulated lines were found to have significantly higher extractability when exposed to NaOH base extraction, indicating altered cell wall construction. The molecular weight of isolated lignin was measured and lignin structure was determined by HSQC NMR-based lignin subunit analysis for control and themore » C3'H and C4H downregulated lines. The slight reductions in average molecular weights of the lignin isolated from the transgenic lines (C3'H = 7000, C4H = 6500, control = 7300) does not appear to explain the difference in extractability. The HSQC NMR-based lignin subunit analysis showed increases in H lignin content for the C3'H but only slight differences in the lignin subunit structure of the C3'H and C4H downregulated lines when compared to the control. The greatest difference between the C3'H and C4H downregulated lines is the total lignin content; therefore, it appears that overall lowered lignin content contributes greatly to reduced recalcitrance and increased extractability of cell wall biopolymers. Furthermore, studies will be conducted to determine how the reduction in lignin content creates a less rigid cell wall that is more prone to extraction and sugar release.« less

Initial Expansion of C4 Vegetation in Australia During the Late Pliocene

NASA Astrophysics Data System (ADS)

Andrae, J. W.; McInerney, F. A.; Polissar, P. J.; Sniderman, J. M. K.; Howard, S.; Hall, P. A.; Phelps, S. R.

2018-05-01

Since the late Miocene, plants using the C4 photosynthetic pathway have increased to become major components of many tropical and subtropical ecosystems. However, the drivers for this expansion remain under debate, in part because of the varied histories of C4 vegetation on different continents. Australia hosts the highest dominance of C4 vegetation of all continents, but little is known about the history of C4 vegetation there. Carbon isotope ratios of plant waxes from scientific ocean drilling sediments off north-western Australia reveal the onset of Australian C4 expansion at 3.5 Ma, later than in many other regions. Pollen analysis from the same sediments reveals increasingly open C3-dominated biomes preceding the shift to open C4-dominated biomes by several million years. We hypothesize that the development of a summer monsoon climate beginning in the late Pliocene promoted a highly seasonal precipitation regime favorable to the expansion of C4 vegetation.

Yield responses of wild C3 and C4 crop progenitors to subambient CO2 : a test for the role of CO2 limitation in the origin of agriculture.

PubMed

Cunniff, Jennifer; Jones, Glynis; Charles, Michael; Osborne, Colin P

2017-01-01

Limitation of plant productivity by the low partial pressure of atmospheric CO 2 (C a ) experienced during the last glacial period is hypothesized to have been an important constraint on the origins of agriculture. In support of this hypothesis, previous work has shown that glacial C a limits vegetative growth in the wild progenitors of both C 3 and C 4 founder crops. Here, we present data showing that glacial C a also reduces grain yield in both crop types. We grew four wild progenitors of C 3 (einkorn wheat and barley) and C 4 crops (foxtail and broomcorn millets) at glacial and postglacial C a , measuring grain yield and the morphological and physiological components contributing to these yield changes. The C 3 species showed a significant increase in unthreshed grain yield of ~50% with the glacial to postglacial increase in C a , which matched the stimulation of photosynthesis, suggesting that increases in photosynthesis are directly translated into yield at subambient levels of C a . Increased yield was controlled by a higher rate of tillering, leading to a larger number of tillers bearing fertile spikes, and increases in seed number and size. The C 4 species showed smaller, but significant, increases in grain yield of 10-15%, arising from larger seed numbers and sizes. Photosynthesis was enhanced by C a in only one C 4 species and the effect diminished during development, suggesting that an indirect mechanism mediated by plant water relations could also be playing a role in the yield increase. Interestingly, the C 4 species at glacial C a showed some evidence that photosynthetic capacity was upregulated to enhance carbon capture. Development under glacial C a also impacted negatively on the subsequent germination and viability of seeds. These results suggest that the grain production of both C 3 and C 4 crop progenitors was limited by the atmospheric conditions of the last glacial period, with important implications for the origins of agriculture. © 2016

Preparation of WO3/g-C3N4 composites and their application in oxidative desulfurization

NASA Astrophysics Data System (ADS)

Zhao, Rongxiang; Li, Xiuping; Su, Jianxun; Gao, Xiaohan

2017-01-01

WO3/graphitic carbon nitride (g-C3N4) composites were successfully synthesized through direct calcining of a mixture of WO3 and g-C3N4 at 400 °C for 2 h. The WO3 was prepared by calcination of phosphotungstic acid at 550 °C for 4 h, and the g-C3N4 was obtained by calcination of melamine at 520 °C for 4 h. The WO3/g-C3N4 composites were characterized by X-ray diffraction (XRD), Scanning electron microscopy (SEM), Fourier-transform infrared spectroscopy (FT-IR), and Brunner-Emmett-Teller analysis (BET). The WO3/g-C3N4 composites exhibited stronger XRD peaks of WO3 and g-C3N4 than the WO3 and pure g-C3N4. In addition, two WO3 peaks at 25.7° and 26.6° emerged for the 36% -WO3/g-C3N4 composite. This finding indicated that WO3 was highly dispersed on the surface of the g-C3N4 nanosheets and interacted with the nanosheets, which resulted in the appearance of (012) and (022) planes of WO3. The WO3/g-C3N4 composite also exhibited a larger specific surface area and higher degree of crystallization than WO3 or pure g-C3N4, which resulted in high catalytic activity of the catalyst. Desulfurization experiments demonstrated that the desulfurization rate of dibenzothiophene (DBT) in model oil reached 91.2% under optimal conditions. Moreover, the activity of the catalyst was not significantly decreased after five recycles.

Synthetic, Infrared, 1H and 13C NMR Spectral Studies on N-(2-/3-Substituted Phenyl)-4-Substituted Benzenesulphonamides, 4-X'C6H4SO2NH(2-/3-XC6H4), where X' = H, CH3, C2H5, F, Cl or Br, and X = CH3 or Cl

NASA Astrophysics Data System (ADS)

Gowda, B. Thimme; Shetty, Mahesha; Jayalakshmi, K. L.

2005-02-01

Twenty three N-(2-/3-substituted phenyl)-4-substituted benzenesulphonamides of the general formula, 4-X'C6H4SO2NH(2-/3-XC6H4), where X' = H, CH3, C2H5, F, Cl or Br and X = CH3 or Cl have been prepared and characterized, and their infrared spectra in the solid state, 1H and 13C NMR spectra in solution were studied. The N-H stretching vibrations, νN-H, absorb in the range 3285 - 3199 cm-1, while the asymmetric and symmetric SO2 vibrations vary in the ranges 1376 - 1309 cm-1 and 1177 - 1148 cm-1, respectively. The S-N and C-N stretching vibrations absorb in the ranges 945 - 893 cm-1 and 1304 - 1168 cm-1, respectively. The compounds do not exhibit particular trends in the variation of these frequencies on substitution either at ortho or meta positions with either a methyl group or Cl. The observed 1H and 13C chemical shifts of are assigned to protons and carbons of the two benzene rings. Incremental shifts of the ring protons and carbons due to -SO2NH(2-/3-XC6H4) groups in C6H5SO2NH(2-/3-XC6H4), and 4- X'C6H4SO2- and 4-X'C6H4SO2NH- groups in 4-X'C6H4SO2NH(C6H5) are computed and employed to calculate the chemical shifts of the ring protons and carbons in the substituted compounds, 4-X'C6H4SO2NH(2-/3-XC6H4). The computed values agree well with the observed chemical shifts.

Step Free Surface Heteroepitaxy of 3C-SiC Layers on Patterned 4H/6H-SiC Mesas and Cantilevers

NASA Technical Reports Server (NTRS)

Neudeck, P. G.; Powell, J. A.; Trunek, A. J.; Spry, D. J.

2004-01-01

The off-axis approach to SiC epitaxial growth has not prevented many substrate crystal defects from propagating into SiC epilayers, and does not permit the realization of SiC heteropolytype devices. This paper reviews recent advancements in SiC epitaxial growth that begin to overcome the above shortcomings for arrays of device-sized mesas patterned into on-axis 4H/6HSiC wafers. These on-axis mesa growth techniques have produced 4H/6H-SiC homoepilayers and 3C-SiC heteroepilayers with substantially lower dislocation densities. The results should enable improved homojunction and heterojunction silicon carbide prototype devices.

Kinetics of the R + NO2 reactions (R = i-C3H7, n-C3H7, s-C4H9, and t-C4H9) in the temperature range 201-489 K.

PubMed

Rissanen, Matti P; Arppe, Suula L; Eskola, Arkke J; Tammi, Matti M; Timonen, Raimo S

2010-04-15

The bimolecular rate coefficients of four alkyl radical reactions with NO(2) have been measured in direct time-resolved experiments. Reactions were studied under pseudo-first-order conditions in a temperature-controlled tubular flow reactor coupled to a laser photolysis/photoionization mass spectrometer (LP-PIMS). The measured reaction rate coefficients are independent of helium bath gas pressure within the experimental ranges covered and exhibit negative temperature dependence. For i-C(3)H(7) + NO(2) and t-C(4)H(9) + NO(2) reactions, the dependence of ordinate (logarithm of reaction rate coefficients) on abscissa (1/T or log(T)) was nonlinear. The obtained results (in cm(3) s(-1)) can be expressed by the following equations: k(n-C(3)H(7) + NO(2)) = ((4.34 +/- 0.08) x 10(-11)) (T/300 K)(-0.14+/-0.08) (203-473 K, 1-7 Torr), k(i-C(3)H(7) + NO(2)) = ((3.66 +/- 2.54) x 10(-12)) exp(656 +/- 201 K/T)(T/300 K)(1.26+/-0.68) (220-489 K, 1-11 Torr), k(s-C(4)H(9) + NO(2)) = ((4.99 +/- 0.16) x 10(-11))(T/300 K)(-1.74+/-0.12) (241-485 K, 2 - 12 Torr) and k(t-C(4)H(9) + NO(2)) = ((8.64 +/- 4.61) x 10(-12)) exp(413 +/- 154 K/T)(T/300 K)(0.51+/-0.55) (201-480 K, 2-11 Torr), where the uncertainties shown refer only to the 1 standard deviations obtained from the fitting procedure. The estimated overall uncertainty in the determined bimolecular rate coefficients is about +/-20%.

Aluminum acceptor four particle bound exciton complex in 4H, 6H, and 3C SiC

NASA Technical Reports Server (NTRS)

Clemen, L. L.; Devaty, R. P.; Macmillan, M. F.; Yoganathan, M.; Choyke, W. J.; Larkin, D. J.; Powell, J. A.; Edmond, J. A.; Kong, H. S.

1993-01-01

Evidence is presented for a four particle acceptor complex in 3C, 6H, and 4H SiC, obtained in low-temperature photoluminescence and cathodoluminescence experiments. The new lines were observed in p-type films lightly doped with aluminum, of 6H, 4H, and 3C SiC grown on the silicon (0001) face of 6H SiC under special conditions. The lines increase in intensity as more aluminum is added during growth. The multiplicity of observed lines is consistent with symmetry-based models similar to those which have been proposed to describe 4A centers in p-type zincblende semiconductors.

Biogenic Volatile Organic Compound and Respiratory CO2 Emissions after 13C-Labeling: Online Tracing of C Translocation Dynamics in Poplar Plants

PubMed Central

Ghirardo, Andrea; Gutknecht, Jessica; Zimmer, Ina; Brüggemann, Nicolas; Schnitzler, Jörg-Peter

2011-01-01

Background Globally plants are the primary sink of atmospheric CO2, but are also the major contributor of a large spectrum of atmospheric reactive hydrocarbons such as terpenes (e.g. isoprene) and other biogenic volatile organic compounds (BVOC). The prediction of plant carbon (C) uptake and atmospheric oxidation capacity are crucial to define the trajectory and consequences of global environmental changes. To achieve this, the biosynthesis of BVOC and the dynamics of C allocation and translocation in both plants and ecosystems are important. Methodology We combined tunable diode laser absorption spectrometry (TDLAS) and proton transfer reaction mass spectrometry (PTR-MS) for studying isoprene biosynthesis and following C fluxes within grey poplar (Populus x canescens) saplings. This was achieved by feeding either 13CO2 to leaves or 13C-glucose to shoots via xylem uptake. The translocation of 13CO2 from the source to other plant parts could be traced by 13C-labeled isoprene and respiratory 13CO2 emission. Principal Finding In intact plants, assimilated 13CO2 was rapidly translocated via the phloem to the roots within 1 hour, with an average phloem transport velocity of 20.3±2.5 cm h−1. 13C label was stored in the roots and partially reallocated to the plants' apical part one day after labeling, particularly in the absence of photosynthesis. The daily C loss as BVOC ranged between 1.6% in mature leaves and 7.0% in young leaves. Non-isoprene BVOC accounted under light conditions for half of the BVOC C loss in young leaves and one-third in mature leaves. The C loss as isoprene originated mainly (76–78%) from recently fixed CO2, to a minor extent from xylem-transported sugars (7–11%) and from photosynthetic intermediates with slower turnover rates (8–11%). Conclusion We quantified the plants' C loss as respiratory CO2 and BVOC emissions, allowing in tandem with metabolic analysis to deepen our understanding of ecosystem C flux. PMID:21387007

Potential individual versus simultaneous climate change effects on soybean (C 3) and maize (C 4) crops: An agrotechnology model based study

NASA Astrophysics Data System (ADS)

Mera, Roberto J.; Niyogi, Dev; Buol, Gregory S.; Wilkerson, Gail G.; Semazzi, Fredrick H. M.

2006-11-01

Landuse/landcover change induced effects on regional weather and climate patterns and the associated plant response or agricultural productivity are coupled processes. Some of the basic responses to climate change can be detected via changes in radiation ( R), precipitation ( P), and temperature ( T). Past studies indicate that each of these three variables can affect LCLUC response and the agricultural productivity. This study seeks to address the following question: What is the effect of individual versus simultaneous changes in R, P, and T on plant response such as crop yields in a C 3 and a C 4 plant? This question is addressed by conducting model experiments for soybean (C 3) and maize (C 4) crops using the DSSAT: Decision Support System for Agrotechnology Transfer, CROPGRO (soybean), and CERES-Maize (maize) models. These models were configured over an agricultural experiment station in Clayton, NC [35.65°N, 78.5°W]. Observed weather and field conditions corresponding to 1998 were used as the control. In the first set of experiments, the CROPGRO (soybean) and CERES-Maize (maize) responses to individual changes in R and P (25%, 50%, 75%, 150%) and T (± 1, ± 2 °C) with respect to control were studied. In the second set, R, P, and T were simultaneously changed by 50%, 150%, and ± 2 °C, and the interactions and direct effects of individual versus simultaneous variable changes were analyzed. For the model setting and the prescribed environmental changes, results from the first set of experiments indicate: (i) precipitation changes were most sensitive and directly affected yield and water loss due to evapotranspiration; (ii) radiation changes had a non-linear effect and were not as prominent as precipitation changes; (iii) temperature had a limited impact and the response was non-linear; (iv) soybeans and maize responded differently for R, P, and T, with maize being more sensitive. The results from the second set of experiments indicate that simultaneous

Isoelectric point and adsorption activity of porous g-C3N4

NASA Astrophysics Data System (ADS)

Zhu, Bicheng; Xia, Pengfei; Ho, Wingkei; Yu, Jiaguo

2015-07-01

The isoelectric point (IEP) is an important physicochemical parameter of many compounds, such as oxides, hydroxides, and nitrides, and can contribute to estimation of the surface charges of compound particles at various pH conditions. In this work, three types of graphitic carbon nitrides (g-C3N4) were synthesized by directly heating melamine, thiourea, and urea. The prepared samples showed different microstructures and IEPs that influenced their adsorption activity. Differences in microstructure resulted from the various precursors used during synthesis. The IEPs of the obtained g-C3N4 were measured to be approximately 4-5, which is due to the equilibrium of chemical reactions between hydrogen ions, hydroxyl ions, and amine groups on the g-C3N4 surface. The IEP of g-C3N4 prepared from thiourea was lower than those of the corresponding samples prepared from melamine and urea. The adsorption activity of methylene blue on g-C3N4 prepared from urea and thiourea was excellent, which indicates that g-C3N4 is a promising adsorbent. This work provides a useful reference for choosing precursors with which to prepare g-C3N4 and combining g-C3N4 with other compounds in solution.