Oxygen Requirement and Inhibition of C4 Photosynthesis1 : An Analysis of C4 Plants Deficient in the C3 and C4 Cycles

The basis for O2 sensitivity of C4 photosynthesis was evaluated using a C4-cycle-limited mutant of Amaranthus edulis (a phosphoenolpyruvate carboxylase-deficient mutant), and a C3-cycle-limited transformant of Flaveria bidentis (an antisense ribulose-1,5-bisphosphate carboxylase/oxygenase [Rubisco] small subunit transformant). Data obtained with the C4-cycle-limited mutant showed that atmospheric levels of O2 (20 kPa) caused increased inhibition of photosynthesis as a result of higher levels of photorespiration. The optimal O2 partial pressure for photosynthesis was reduced from approximately 5 kPa O2 to 1 to 2 kPa O2, becoming similar to that of C3 plants. Therefore, the higher O2 requirement for optimal C4 photosynthesis is specifically associated with the C4 function. With the Rubisco-limited F. bidentis, there was less inhibition of photosynthesis by supraoptimal levels of O2 than in the wild type. When CO2 fixation by Rubisco is limited, an increase in the CO2 concentration in bundle-sheath cells via the C4 cycle may further reduce the oxygenase activity of Rubisco and decrease the inhibition of photosynthesis by high partial pressures of O2 while increasing CO2 leakage and overcycling of the C4 pathway. These results indicate that in C4 plants the investment in the C3 and C4 cycles must be balanced for maximum efficiency.

Nitrogen washing from C3 and C4 cover grasses residues by rain

Crop species with the C-4 photosynthetic pathway are more efficient in assimilating N than C-3 plants, which results in different N amounts prone to be washed from its straw by rain water. Such differences may affect N recycling in agricultural systems where these species are grown as cover crops. In this experiment, phytomass production and N leaching from the straw of grasses with different photosynthetic pathways were studied in response to N application. Pearl millet (Pennisetum glaucum) and congo grass (Brachiaria ruziziensis) with the C-4 photosynthetic pathway, and black oat (Arena Strigosa) and triticale (X Triticosecale), with the C-3 photosynthetic pathway, were grown for 47 days. After determining dry matter yields and N and C contents, a 30 mm rainfall was simulated over 8 t ha(-1) of dry matter of each plant residue and the leached amounts of ammonium and nitrate were determined. C-4 grasses responded to higher fertilizer rates, whereas N contents in plant tissue were lower. The amount of N leached from C-4 grass residues was lower, probably because the C/N ratio is higher and N is more tightly bound to organic compounds. When planning a crop rotation system it is important to take into account the difference in N release of different plant residues which may affect N nutrition of the subsequent crop.

Carbon isotope composition as a tool for quality control of herbs, spices and medicinal plants

Isotope screening is a simple and cheap test for determining the photosynthetic pathway used by plants. The scope of this work was to classify the photosynthetic type of Melissa officinalis L. and Cymbopogon citratus [DC.] Stapf, through studies of the carbon isotope composition (δ13 C), and we are proposing the use of carbon isotope composition results as a tool to control the quality of medicinal plants. For studies of δ 13C (13 C% = [R (sample)/R (standard) - 1] × 10 -3), dried, powdered leaves were analyzed in a mass spectrometer coupled with an elemental analyzer for determining the ratio R (R = 13CO2/12CO2). As results, M. officinalis presented a C3 photosynthetic type, and C. citratus presented a C4 photosynthetic type. The carbon isotope composition from this study can be used as quality control of M. officinalis adulterants.

Estimating Photosynthesis and Concurrent Export Rates in C3 and C4 Species at Ambient and Elevated CO21 2

The ability of 21 C3 and C4 monocot and dicot species to rapidly export newly fixed C in the light at both ambient and enriched CO2 levels was compared. Photosynthesis and concurrent export rates were estimated during isotopic equilibrium of the transport sugars using a steady-state 14CO2-labeling procedure. At ambient CO2 photosynthesis and export rates for C3 species were 5 to 15 and 1 to 10 μmol C m−2 s−1, respectively, and 20 to 30 and 15 to 22 μmol C m−2 s−1, respectively, for C4 species. A linear regression plot of export on photosynthesis rate of all species had a correlation coefficient of 0.87. When concurrent export was expressed as a percentage of photosynthesis, several C3 dicots that produced transport sugars other than Suc had high efflux rates relative to photosynthesis, comparable to those of C4 species. At high CO2 photosynthetic and export rates were only slightly altered in C4 species, and photosynthesis increased but export rates did not in all C3 species. The C3 species that had high efflux rates relative to photosynthesis at ambient CO2 exported at rates comparable to those of C4 species on both an absolute basis and as a percentage of photosynthesis. At ambient CO2 there were strong linear relationships between photosynthesis, sugar synthesis, and concurrent export. However, at high CO2 the relationships between photosynthesis and export rate and between sugar synthesis and export rate were not as strong because sugars and starch were accumulated.

Comparison of Belowground Biomass in C3- and C4-Dominated Mixed Communities in a Chesapeake Bay Brackish Marsh

Belowground biomass is a critical factor regulating ecosystem functions of coastal marshes, including soil organic matter (SOM) accumulation and the ability of these systems to keep pace with sea-level rise. Nevertheless, belowground biomass responses to environmental and vegetation changes have been given little emphasis marsh studies. Here we present a method using stable carbon isotopes and color to identify root and rhizomes of Schoenoplectus americanus (Pers.) Volk. ex Schinz and R. Keller (C3) and Spartina patens (Ait.) Muhl. (C4) occurring in C3− and C4-dominated communities in a Chesapeake Bay brackish marsh. The functional significance of the biomass classes we identified is underscored by differences in their chemistry, depth profiles, and variation in biomass and profiles relative to abiotic and biotic factors. C3 rhizomes had the lowest concentrations of cellulose (29.19%) and lignin (14.43%) and the lowest C:N (46.97) and lignin:N (0.16) ratios. We distinguished two types of C3 roots, and of these, the dark red C3 roots had anomalously high C:N (195.35) and lignin:N (1.14) ratios, compared with other root and rhizome classes examined here and with previously published values. The C4-dominated community had significantly greater belowground biomass (4119.1 g m−2) than the C3-dominated community (3256.9 g m−2), due to greater total root biomass and a 3.6-fold higher C3-root:rhizome ratio in the C4-dominated community. C3 rhizomes were distributed significantly shallower in the C4-dominated community, while C3 roots were significantly deeper. Variability in C3 rhizome depth distributions was explained primarily by C4 biomass, and C3 roots were explained primarily by water table height. Our results suggest that belowground biomass in this system is sensitive to slight variations in water table height (across an 8 cm range), and that the reduced overlap between C3 and C4 root profiles in the C4-dominated community may account for the greater total root biomass observed in that community. Given that future elevated atmospheric CO2 and accelerated sea-level rise are likely to increase C3 abundance in Atlantic and Gulf coast marshes, investigations that quantify how patterns of C3 and C4 belowground biomass respond to environmental and biological factors stand to improve our understanding of ecosystem-wide impacts of global changes on coastal wetlands.

Caracterização sistêmica de alterações em redes fisiológicas de plantas C3 e C4 submetidas à deficiência hídrica

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Avaliação metabólica de dietas de C3 e C4 na formação do ovo (gema e albúmen) pelo uso da técnica dos isótopos estáveis do carbono

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Comparison between the C5 or C6-Cz electrode assembly and C3 or C4-Cz assembly for transcranial electric motor activation of muscular response of the contralateral facial nerve

We used an assembly of electrodes C3 and C4-Cz in order to activate the motor cortical area of the corticobulbar tract to elucidate the motor-evoked potential of the contralateral mentalis muscle. We compared this setup to that of an assembly with electrodes C5 or C6-Cz using a train of electrical pulses and a single electrical pulse. This analysis was made in 23 consecutive patients who underwent several varied surgeries and were prospectively operated on at Santa Paula Hospital between January and June 2011. The results showed that the assembly with C5 or C6-Cz produced a multisynaptic motor-evoked potential in the contralateral mentalis muscle in 86.9 % of the patients, whereas 82.6 % of patients stimulated at points C3 or C4-Cz presented the same response. However, both assemblies showed similar behavior with the use of a single electrical pulse for peripheral contralateral nerve stimulation. We concluded that the C5 or C6-Cz assembly was similar to C3 or C4-Cz in obtaining a multisynaptic response in the contralateral mentalis muscle, although it required less intensive stimulation than the C3 or C4- Cz assembly.

n-alkane carbon isotope ratios in plants and recent lake sediments from Cameroon

Trees and shrubs in tropical Africa use the C3 cycle as a carbon fixation pathway during photosynthesis, while grasses and sedges mostly use the C4 cycle. Leaf-wax lipids from sedimentary archives such as the long-chain n-alkanes (e.g., n-C27 to n-C33) inherit carbon isotope ratios that are representative of the carbon fixation pathway. Therefore, n-alkane d13C values are often used to reconstruct past C3/C4 composition of vegetation, assuming that the relative proportions of C3 and C4 leaf waxes reflect the relative proportions of C3 and C4 plants. We have compared the d13C values of n-alkanes from modern C3 and C4 plants with previously published values from recent lake sediments and provide a framework for estimating the fractional contribution (areal-based) of C3 vegetation cover (fC3) represented by these sedimentary archives. Samples were collected in Cameroon, across a latitudinal transect that accommodates a wide range of climate zones and vegetation types, as reflected in the progressive northward replacement of C3-dominated rain forest by C4-dominated savanna. The C3 plants analysed were characterised by substantially higher abundances of n-C29 alkanes and by substantially lower abundances of n-C33 alkanes than the C4 plants. Furthermore, the sedimentary d13C values of n-C29 and n-C31 alkanes from recent lake sediments in Cameroon (-37.4 per mil to -26.5 per mil) were generally within the range of d13C values for C3 plants, even when from sites where C4 plants dominated the catchment vegetation. In such cases simple linear mixing models fail to accurately reconstruct the relative proportions of C3 and C4 vegetation cover when using the d13C values of sedimentary n-alkanes, overestimating the proportion of C3 vegetation, likely as a consequence of the differences in plant wax production, preservation, transport, and/or deposition between C3 and C4 plants. We therefore tested a set of non-linear binary mixing models using d13C values from both C3 and C4 vegetation as end-members. The non-linear models included a sigmoid function (sine-squared) that describes small variations in the fC3 values as the minimum and maximum d13C values are approached, and a hyperbolic function that takes into account the differences between C3 and C4 plants discussed above. Model fitting and the estimation of uncertainties were completed using the Monte Carlo algorithm and can be improved by future data addition. Models that provided the best fit with the observed d13C values of sedimentary n-alkanes were either hyperbolic functions or a combination of hyperbolic and sine-squared functions. Such non-linear models may be used to convert d13C measurements on sedimentary n-alkanes directly into reconstructions of C3 vegetation cover.

Understanding 2H/1H systematics of leaf wax n-alkanes in coastal plants at Stiffkey saltmarsh, Norfolk, UK

Interpretation of sedimentary n-alkyl lipid d2H data is complicated by a limited understanding of factors controlling interspecies variation in biomarker 2H/1H composition. To distinguish between the effects of interrelated environmental, physical and biochemical controls on the hydrogen isotope composition of n-alkyl lipids, we conducted linked d2H analyses of soil water, xylem water, leaf water and n-alkanes from a range of C3 and C4 plants growing at a UK saltmarsh (i) across multiple sampling sites, (ii) throughout the 2012 growing season, and (iii) at different times of the day. Soil waters varied isotopically by up to 35& depending on marsh sub-environment, and exhibited site-specific seasonal shifts in d2H up to a maximum of 31 per mil. Maximum interspecies variation in xylem water was 38 per mil, while leaf waters differed seasonally by a maximum of 29 per mil. Leaf wax n-alkane 2H/1H, however, consistently varied by over 100 per mil throughout the 2012 growing season, resulting in an interspecies range in the ewax/leaf water values of -79 per mil to –227 per mil. From the discrepancy in the magnitude of these isotopic differences, we conclude that mechanisms driving variation in the 2H/1H composition of leaf water, including (i) spatial changes in soil water 2H/1H, (ii) temporal changes in soil water 2H/1H, (iii) differences in xylem water 2H/1H, and (iv) differences in leaf water evaporative 2H-enrichment due to varied plant life forms, cannot explain the range of n-alkane d2H values we observed. Results from this study suggests that accurate reconstructions of palaeoclimate regimes from sedimentary n-alkane d2H require further research to constrain those biological mechanisms influencing species-specific differences in 2H/1H fractionation during lipid biosynthesis, in particular where plants have developed biochemical adaptations to water-stressed conditions. Understanding how these mechanisms interact with environmental conditions will be crucial to ensure accurate interpretation of hydrogen isotope signals from the geological record.