SciCombinator

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Concept: Carbon fixation

165

Agaves are succulent monocotyledonous plants native to xeric environments of North America. Because of their adaptations to their environment, including crassulacean acid metabolism (CAM, a water-efficient form of photosynthesis), and existing technologies for ethanol production, agaves have gained attention both as potential lignocellulosic bioenergy feedstocks and models for exploring plant responses to abiotic stress. However, the lack of comprehensive Agave sequence datasets limits the scope of investigations into the molecular-genetic basis of Agave traits.

Concepts: Photosynthesis, C4 carbon fixation, Crassulacean acid metabolism, Agave, Carbon fixation, Agave tequilana

165

Crassulacean acid metabolism (CAM) occurs in many Euphorbiaceae, particularly Euphorbia, a genus with C3 and C4 species as well. With the aim of contributing to our knowledge of the evolution of CAM in this genus, this study examined the possible occurrence of CAM in Euphorbia milii, a species with leaf succulence and drought tolerance suggestive of this carbon fixation pathway. Leaf anatomy consisted of a palisade parenchyma, a spongy parenchyma and a bundle sheath with chloroplasts, which indicates the possible functioning of C2 photosynthesis. No evidence of nocturnal CO2 fixation was found in plants of E. milii either watered or under drought; watered plants had a low nocturnal respiration rate ®. After 12 days without watering, the photosynthetic rate (P N) decreased 85 % and nocturnal R was nearly zero. Nocturnal H(+) accumulation (ΔH(+)) in watered plants was 18 ± 2 (corresponding to malate) and 18 ± 4 (citrate) μmol H(+) (g fresh mass)(-1). Respiratory CO2 recycling through acid synthesis contributed to a night-time water saving of 2 and 86 % in watered plants and plants under drought, respectively. Carbon isotopic composition (δ(13)C) was -25.2 ± 0.7 ‰ in leaves and -24.7 ± 0.1 ‰ in stems. Evidence was found for the operation of weak CAM in E. milii, with statistically significant ΔH(+), no nocturnal CO2 uptake and values of δ(13)C intermediate between C3 and constitutive CAM plants; ΔH(+) was apparently attributable to both malate and citrate. The results suggest that daily malate accumulation results from recycling of part of the nocturnal respiratory CO2, which helps explain the occurrence of an intermediate value of leaf δ(13)C. Euphorbia milii can be considered as a CAM-cycling species. The significance of the operation of CAM-cycling in E. milii lies in water conservation, rather than carbon acquisition. The possible occurrence of C2 photosynthesis merits research.

Concepts: Photosynthesis, Oxygen, Carbon dioxide, Water, C4 carbon fixation, C3 carbon fixation, Crassulacean acid metabolism, Carbon fixation

162

In several taxa, increasing leaf succulence has been associated with decreasing mesophyll conductance (g M) and an increasing dependence on Crassulacean acid metabolism (CAM). However, in succulent Aizoaceae, the photosynthetic tissues are adjacent to the leaf surfaces with an internal achlorophyllous hydrenchyma. It was hypothesized that this arrangement increases g M, obviating a strong dependence on CAM, while the hydrenchyma stores water and nutrients, both of which would only be sporadically available in highly episodic environments. These predictions were tested with species from the Aizoaceae with a 5-fold variation in leaf succulence. It was shown that g M values, derived from the response of photosynthesis to intercellular CO2 concentration (A:C i), were independent of succulence, and that foliar photosynthate δ(13)C values were typical of C3, but not CAM photosynthesis. Under water stress, the degree of leaf succulence was positively correlated with an increasing ability to buffer photosynthetic capacity over several hours and to maintain light reaction integrity over several days. This was associated with decreased rates of water loss, rather than tolerance of lower leaf water contents. Additionally, the hydrenchyma contained ~26% of the leaf nitrogen content, possibly providing a nutrient reservoir. Thus the intermittent use of C3 photosynthesis interspersed with periods of no positive carbon assimilation is an alternative strategy to CAM for succulent taxa (Crassulaceae and Aizoaceae) which occur sympatrically in the Cape Floristic Region of South Africa.

Concepts: Photosynthesis, Carbon dioxide, Leaf, RuBisCO, C4 carbon fixation, C3 carbon fixation, Crassulacean acid metabolism, Carbon fixation

28

Background and AimsA positive correlation between tissue thickness and crassulacean acid metabolism (CAM) expression has been frequently suggested. Therefore, this study addressed the question of whether water availability modulates photosynthetic plasticity in different organs of two epiphytic orchids with distinct leaf thickness.MethodsTissue morphology and photosynthetic mode (C3 and/or CAM) were examined in leaves, pseudobulbs and roots of a thick-leaved (Cattleya walkeriana) and a thin-leaved (Oncidium ‘Aloha’) epiphytic orchid. Morphological features were studied comparing the drought-induced physiological responses observed in each organ after 30 d of either drought or well-watered treatments.Key ResultsCattleya walkeriana, which is considered a constitutive CAM orchid, displayed a clear drought-induced up-regulation of CAM in its thick leaves but not in its non-leaf organs (pseudobulbs and roots). The set of morphological traits of Cattleya leaves suggested the drought-inducible CAM up-regulation as a possible mechanism of increasing water-use efficiency and carbon economy. Conversely, although belonging to an orchid genus classically considered as performing C3 photosynthesis, Oncidium ‘Aloha’ under drought seemed to express facultative CAM in its roots and pseudobulbs but not in its leaves, indicating that such photosynthetic responses might compensate for the lack of capacity to perform CAM in its thin leaves. Morphological features of Oncidium leaves also indicated lower efficiency in preventing water and CO2 losses, while aerenchyma ducts connecting pseudobulbs and leaves suggested a compartmentalized mechanism of nighttime carboxylation via phosphoenolpyruvate carboxylase (PEPC) (pseudobulbs) and daytime carboxylation via Rubisco (leaves) in drought-exposed Oncidium plants.ConclusionsWater availability modulated CAM expression in an organ-compartmented manner in both orchids studied. As distinct regions of the same orchid could perform different photosynthetic pathways and variable degrees of CAM expression depending on the water availability, more attention should be addressed to this in future studies concerning the abundance of CAM plants.

Concepts: Photosynthesis, Carbon dioxide, RuBisCO, C4 carbon fixation, C3 carbon fixation, Crassulacean acid metabolism, Orchidaceae, Carbon fixation

28

The activity of the photosynthetic carbon-fixing enzyme, ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco), is partially inhibited by arsenite in the millimolar concentration range. However, micromolar arsenite can fully inhibit Rubisco in the presence of a potentiating monothiol such as cysteine, cysteamine, 2-mercaptoethanol or N-acetylcysteine, but not glutathione. Arsenite reacts specifically with the vicinal Cys172-Cys192 from the large subunit of Rubisco and with the monothiol to establish a ternary complex which is suggested to be a trithioarsenical. The stability of the complex is strongly dependent on the nature of the monothiol. Enzyme activity is fully recovered through the disassembly of the complex after eliminating arsenite and/or the thiol from the medium. The synergic combination of arsenite and a monothiol acts also in vivo stopping carbon dioxide fixation in illuminated cultures of Chlamydomonas reinhardtii. Again, this effect may be reverted by washing the cells. However, in vivo inhibition does not result from the blocking of Rubisco since mutant strains carrying Rubiscos with Cys172 and/or Cys192 substitutions (which are insensitive to arsenite in vitro) are also arrested. This suggests the existence of a specific sensor controlling carbon fixation that is even more sensitive than Rubisco to the arsenite-thiol synergism.

Concepts: Photosynthesis, Oxygen, Carbon dioxide, Enzyme inhibitor, Glutathione, Calvin cycle, Thiol, Carbon fixation

27

Stable carbon isotope ratios (δ(13) C) of terrestrial plants are employed across a diverse range of applications in environmental and plant sciences; however, the kind of information that is desired from the δ(13) C signal often differs. At the extremes, it ranges between purely environmental and purely biological. Here, we review environmental drivers of variation in carbon isotope discrimination (Δ) in terrestrial plants, and the biological processes that can either damp or amplify the response. For C3 plants, where Δ is primarily controlled by the ratio of intercellular to ambient CO2 concentrations (ci /ca ), coordination between stomatal conductance and photosynthesis and leaf area adjustment tends to constrain the potential environmentally driven range of Δ. For C4 plants, variation in bundle-sheath leakiness to CO2 can either damp or amplify the effects of ci /ca on Δ. For plants with crassulacean acid metabolism (CAM), Δ varies over a relatively large range as a function of the proportion of daytime to night-time CO2 fixation. This range can be substantially broadened by environmental effects on Δ when carbon uptake takes place primarily during the day. The effective use of Δ across its full range of applications will require a holistic view of the interplay between environmental control and physiological modulation of the environmental signal.

Concepts: Photosynthesis, Carbon dioxide, Ratio, RuBisCO, C4 carbon fixation, C3 carbon fixation, Crassulacean acid metabolism, Carbon fixation

27

Crassulacean acid metabolism (CAM) is a physiological adaptation of plants that live in stress environment conditions. A good model of CAM modulation is the epiphytic bromeliad, Guzmania monostachia, which switches between two photosynthetic pathways (C3-CAM) in response to different environmental conditions, such as light stress and water availability. Along the leaf length a gradient of acidity can be observed when G. monostachia plants are kept under water deficiency. Previous studies showed that the apical portions of the leaves present higher expression of CAM, while the basal regions exhibit lower expression of this photosynthetic pathway. The present study has demonstrated that it is possible to induce the CAM pathway in detached leaves of G. monostachia kept under water deficit for 7 d. Also, it was evaluated whether CAM expression can be modulated in detached leaves of Guzmania and whether some spatial separation between NO3(-) reduction and CO2 fixation occurs in basal and apical portions of the leaf. In addition, we analyzed the involvement of endogenous cytokinins (free and ribosylated forms) as possible signal modulating both NO3(-) reduction and CO2 fixation along the leaf blade of this bromeliad. Besides demonstrating a clear spatial and functional separation of carbon and nitrogen metabolism along G. monostachia leaves, the results obtained also indicated a probable negative correlation between endogenous free cytokinins - zeatin (Z) and isopentenyladenine (iP) - concentration and PEPC activity in the apical portions of G. monostachia leaves kept under water deficit. On the other hand, a possible positive correlation between endogenous Z and iP levels and NR activity in basal portions of drought-exposed and control leaves was verified. Together with the observations presented above, results obtained with exogenous cytokinins treatments, strongly suggest that free cytokinins might act as a stimulatory signal involved in NR activity regulation and as a negative regulator of PEPC activity in CAM-induced leaves of G. monostachia during a diel cycle.

Concepts: Photosynthesis, Carbon dioxide, Bromeliaceae, C4 carbon fixation, C3 carbon fixation, Crassulacean acid metabolism, Carbon fixation, Guzmania

23

Experimental elevation of [CO2] around C3 crops in the field has been shown to increase yields by suppressing the Rubisco oxygenase reaction and in turn photorespiration. Bioengineering a cyanobacterial carbon-concentration mechanism (CCM) into C3 crop species provides a potential means of elevating [CO2] at Rubisco, thereby decreasing photorespiration and increasing photosynthetic efficiency and yield. The cyanobacterial CCM is an attractive alternative relative to other CCMs, because its features do not require anatomical changes to leaf tissue. However, potential benefits of engineering the entire CCM into a C3 leaf are unexamined. Here a CO2 and HCO3- diffusion-reaction model is developed to examine how components of the cyanobacterial CCM affect leaf light-saturated CO2 uptake (Asat) and to determine whether a different Rubisco isoform would perform better in a leaf with a cyanobacterial CCM. Results show that addition of carboxysomes without other CCM components substantially decreases Asat, and that the best first step is addition of HCO3- transporters, as a single HCO3- transporter increased modeled Asat by 9%. Addition of all major CCM components increased Asat from 24 to 38 µmol m-2 ¬s-1. Several Rubisco isoforms were compared in the model, and increasing RuBP-regeneration rate will allow for further improvements by using an Rubisco isoform adapted to high [CO2]. Results from field studies that artificially raise [CO2] suggest that this 60% increase in Asat could result in a 36% to 60% increase in yield.

Concepts: Photosynthesis, Carbon dioxide, Calvin cycle, Photorespiration, RuBisCO, C4 carbon fixation, C3 carbon fixation, Carbon fixation

16

Arundo donax has attracted interest as a potential bioenergy crop due to a high apparent productivity. It uses C3 photosynthesis yet appears competitive with C4 grass biomass feedstock’s and grows in warm conditions where C4 species might be expected to be that productive. Despite this there has been no systematic study of leaf photosynthetic properties. This study determines photosynthetic and photorespiratory parameters for leaves in a natural stand of A. donax growing in southern Portugal. We hypothesise that A. donax has a high photosynthetic potential in high and low light, stomatal limitation to be small and intrinsic water use efficiency unusually low. High photosynthetic rates in A. donax resulted from a high capacity for both maximum Rubisco (Vc,max 117 μmol CO2 m(-2) s(-1)) and ribulose-1:5-bisphosphate limited carboxylation rate (Jmax 213 μmol CO2 m(-2) s(-1)) under light-saturated conditions. Maximum quantum yield for light-limited CO2 assimilation was also high relative to other C3 species. Photorespiratory losses were similar to other C3 species under the conditions of measurement (25%), while stomatal limitation was high (0.25) resulting in a high intrinsic water use efficiency. Overall the photosynthetic capacity of A. donax is high compared to other C3 species, and comparable to C4 bioenergy grasses.

Concepts: Photosynthesis, Carbon dioxide, Plant, Poaceae, Calvin cycle, Leaf, Photorespiration, Carbon fixation

14

In C4 species, the major β-carbonic anhydrase (β-CA) localized in the mesophyll cytosol catalyses the hydration of CO2 to HCO3(-), which phosphoenolpyruvate carboxylase uses in the first step of C4 photosynthesis. To address the role of CA in C4 photosynthesis, we generated transgenic Setaria viridis depleted in β-CA. Independent lines were identified with as little as 13% of wild-type CA. No photosynthetic defect was observed in the transformed lines at ambient CO2 partial pressure (pCO2). At low pCO2, a strong correlation between CO2 assimilation rates and CA hydration rates was observed. C(18)O(16)O isotope discrimination was used to estimate the mesophyll conductance to CO2 diffusion from the intercellular air space to the mesophyll cytosol (gm) in control plants, which allowed us to calculate CA activities in the mesophyll cytosol (Cm). This revealed a strong relationship between the initial slope of the response of the CO2 assimilation rate to cytosolic pCO2 (ACm) and cytosolic CA activity. However, the relationship between the initial slope of the response of CO2 assimilation to intercellular pCO2 (ACi) and cytosolic CA activity was curvilinear. This indicated that in S. viridis, mesophyll conductance may be a contributing limiting factor alongside CA activity to CO2 assimilation rates at low pCO2.

Concepts: Photosynthesis, Oxygen, Carbon dioxide, Metabolism, Redox, PH, Calvin cycle, Carbon fixation