Showing papers on "Photosynthesis published in 2012"

The artificial leaf

Abstract: To convert the energy of sunlight into chemical energy, the leaf splits water via the photosynthetic process to produce molecular oxygen and hydrogen, which is in a form of separated protons and electrons. The primary steps of natural photosynthesis involve the absorption of sunlight and its conversion into spatially separated electron–hole pairs. The holes of this wireless current are captured by the oxygen evolving complex (OEC) of photosystem II (PSII) to oxidize water to oxygen. The electrons and protons produced as a byproduct of the OEC reaction are captured by ferrodoxin of photosystem I. With the aid of ferrodoxin–NADP+ reductase, they are used to produce hydrogen in the form of NADPH. For a synthetic material to realize the solar energy conversion function of the leaf, the light-absorbing material must capture a solar photon to generate a wireless current that is harnessed by catalysts, which drive the four electron/hole fuel-forming water-splitting reaction under benign conditions and under 1 su...

Functions of Macronutrients

Abstract: Publisher Summary This chapter focuses on the role played by various macronutrients such as nitrogen (N), sulfur (S), phosphorus (P), magnesium (Mg), calcium (Ca), and potassium (K) in plant metabolism and growth and describes the symptoms of deficiency and toxicity of these macronutrients. N is the most essential element required after carbon, and it plays a central role in plant metabolism as a constituent of proteins, nucleic acids, chlorophyll, coenzymes, phytohormones, and secondary metabolites. When it is taken as ammonium or nitrate, it is assimilated into amino acids either in the roots or shoots and within the plant, it is translocated as nitrate or amino acids. Sulfur is taken up as sulphate and assimilated into S-containing amino acids such as cysteine that are used to synthesize S-containing enzymes and coenzymes as well as secondary compounds such as phytochelatins (detoxification of metals) or aliins and glucosinolates (feeding deterrents). Phosphorus is a structural element in nucleic acids, and as a component of adenosine phosphates, it plays an important role in energy transfer, and it is also essential for transfer of carbohydrates in leaf cells. Magnesium is a component of chlorophyll, and it is required for photosynthesis and protein synthesis. Calcium is important for cell wall and membrane stabilization, osmoregulation, and as second messenger, thereby allowing plants to regulate developmental processes in response to environmental stimuli. The main role of K is osmoregulation, which is important for cell extension and stomata movement, and it affects loading of sucrose and the rate of mass flow-driven solute movement within the plant.

Photorespiration and the Evolution of C4 Photosynthesis

Abstract: C(4) photosynthesis is one of the most convergent evolutionary phenomena in the biological world, with at least 66 independent origins. Evidence from these lineages consistently indicates that the C(4) pathway is the end result of a series of evolutionary modifications to recover photorespired CO(2) in environments where RuBisCO oxygenation is high. Phylogenetically informed research indicates that the repositioning of mitochondria in the bundle sheath is one of the earliest steps in C(4) evolution, as it may establish a single-celled mechanism to scavenge photorespired CO(2) produced in the bundle sheath cells. Elaboration of this mechanism leads to the two-celled photorespiratory concentration mechanism known as C(2) photosynthesis (commonly observed in C(3)-C(4) intermediate species) and then to C(4) photosynthesis following the upregulation of a C(4) metabolic cycle.

Photosynthetic control of electron transport and the regulation of gene expression

Abstract: The term ‘photosynthetic control’ describes the short- and long-term mechanisms that regulate reactions in the photosynthetic electron transport (PET) chain so that the rate of production of ATP and NADPH is coordinated with the rate of their utilization in metabolism. At low irradiances these mechanisms serve to optimize light use efficiency, while at high irradiances they operate to dissipate excess excitation energy as heat. Similarly, the production of ATP and NADPH in ratios tailored to meet demand is finely tuned by a sophisticated series of controls that prevents the accumulation of high NAD(P)H/NAD(P) ratios and ATP/ADP ratios that would lead to potentially harmful over-reduction and inactivation of PET chain components. In recent years, photosynthetic control has also been extrapolated to the regulation of gene expression because mechanisms that are identical or similar to those that serve to regulate electron flow through the PET chain also coordinate the regulated expression of genes encoding photosynthetic proteins. This requires coordinated gene expression in the chloroplasts, mitochondria, and nuclei, involving complex networks of forward and retrograde signalling pathways. Photosynthetic control operates to control photosynthetic gene expression in response to environmental and metabolic changes. Mining literature data on transcriptome profiles of C3 and C4 leaves from plants grown under high atmospheric carbon dioxide (CO2) levels compared with those grown with ambient CO2 reveals that the transition to higher photorespiratory conditions in C3 plants enhances the expression of genes associated with cyclic electron flow pathways in Arabidopsis thaliana, consistent with the higher ATP requirement (relative to NADPH) of photorespiration.

ATP drives direct photosynthetic production of 1-butanol in cyanobacteria

Abstract: While conservation of ATP is often a desirable trait for microbial production of chemicals, we demonstrate that additional consumption of ATP may be beneficial to drive product formation in a nonnatural pathway. Although production of 1-butanol by the fermentative coenzyme A (CoA)-dependent pathway using the reversal of β-oxidation exists in nature and has been demonstrated in various organisms, the first step of the pathway, condensation of two molecules of acetyl-CoA to acetoacetyl-CoA, is thermodynamically unfavorable. Here, we show that artificially engineered ATP consumption through a pathway modification can drive this reaction forward and enables for the first time the direct photosynthetic production of 1-butanol from cyanobacteria Synechococcus elongatus PCC 7942. We further demonstrated that substitution of bifunctional aldehyde/alcohol dehydrogenase (AdhE2) with separate butyraldehyde dehydrogenase (Bldh) and NADPH-dependent alcohol dehydrogenase (YqhD) increased 1-butanol production by 4-fold. These results demonstrated the importance of ATP and cofactor driving forces as a design principle to alter metabolic flux.

Advances in photosynthesis and respiration

Abstract: Photosynthesis in silico: Understanding Complexity from Molecules to Ecosystems is a unique book that aims to show an integrated approach to the understanding of photosynthesis processes. In this volume using mathematical modeling processes are described from the biophysics of the interaction of light with pigment systems to the mutual interaction of individual plants and other organisms in canopies and large ecosystems, up to the global ecosystem issues. Chapters are written by 44 international authorities from 15 countries. Mathematics is a powerful tool for quantitative analysis. Properly programmed, contemporary computers are able to mimic complicated processes in living cells, leaves, canopies and ecosystems. These simulations mathematical models help us predict the photosynthetic responses of modeled systems under various combinations of environmental conditions, potentially occurring in nature, e.g., the responses of plant canopies to globally increasing temperature and atmospheric CO2 concentration. Tremendous analytical power is needed to understand nature's infinite complexity at every level. This book is not a list of equations and computer programs, but the emphasis is on analytical ideas facilitating the understanding of complex interactions governing the photosynthetic process on every level and between different levels of hierarchy. The book provides the necessary background on photosynthesis and demonstrates the benefits of the computer-aided quantitative analysis of its reactions; it is designed for graduate students and researchers in plant physiology, functional plant biology, plant biochemistry, systems biology, biophysics, bio-energy and bio-fuel. more on http://springer.com/978-1-4020-9236-7 2009. Approx. 520 p. ISBN: 978-1-4020-9236-7 ▶ 209,00 €; $279.00; SFr. 347.00; £188.00

The Response of Diatom Central Carbon Metabolism to Nitrogen Starvation Is Different from That of Green Algae and Higher Plants

Abstract: The availability of nitrogen varies greatly in the ocean and limits primary productivity over large areas. Diatoms, a group of phytoplankton that are responsible for about 20% of global carbon fixation, respond rapidly to influxes of nitrate and are highly successful in upwelling regions. Although recent diatom genome projects have highlighted clues to the success of this group, very little is known about their adaptive response to changing environmental conditions. Here, we compare the proteome of the marine diatom Thalassiosira pseudonana (CCMP 1335) at the onset of nitrogen starvation with that of nitrogen-replete cells using two-dimensional gel electrophoresis. In total, 3,310 protein spots were distinguishable, and we identified 42 proteins increasing and 23 decreasing in abundance (greater than 1.5-fold change; P < 0.005). Proteins involved in the metabolism of nitrogen, amino acids, proteins, and carbohydrates, photosynthesis, and chlorophyll biosynthesis were represented. Comparison of our proteomics data with the transcriptome response of this species under similar growth conditions showed good correlation and provided insight into different levels of response. The T. pseudonana response to nitrogen starvation was also compared with that of the higher plant Arabidopsis (Arabidopsis thaliana), the green alga Chlamydomonas reinhardtii, and the cyanobacterium Prochlorococcus marinus. We have found that the response of diatom carbon metabolism to nitrogen starvation is different from that of other photosynthetic eukaryotes and bears closer resemblance to the response of cyanobacteria.

Rerouting carbon flux to enhance photosynthetic productivity.

Abstract: The bioindustrial production of fuels, chemicals, and therapeutics typically relies upon carbohydrate inputs derived from agricultural plants, resulting in the entanglement of food and chemical commodity markets. We demonstrate the efficient production of sucrose from a cyanobacterial species, Synechococcus elongatus, heterologously expressing a symporter of protons and sucrose (cscB). cscB-expressing cyanobacteria export sucrose irreversibly to concentrations of >10 mM without culture toxicity. Moreover, sucrose-exporting cyanobacteria exhibit increased biomass production rates relative to wild-type strains, accompanied by enhanced photosystem II activity, carbon fixation, and chlorophyll content. The genetic modification of sucrose biosynthesis pathways to minimize competing glucose- or sucrose-consuming reactions can further improve sucrose production, allowing the export of sucrose at rates of up to 36.1 mg liter(-1) h illumination(-1). This rate of production exceeds that of previous reports of targeted, photobiological production from microbes. Engineered S. elongatus produces sucrose in sufficient quantities (up to ∼80% of total biomass) such that it may be a viable alternative to sugar synthesis from terrestrial plants, including sugarcane.

Modulation of photosynthetic energy conversion efficiency in nature: from seconds to seasons

Abstract: Modulation of the efficiency with which leaves convert absorbed light to photochemical energy [intrinsic efficiency of open photosystem II (PSII) centers, as the ratio of variable to maximal chlorophyll fluorescence] as well as leaf xanthophyll composition (interconversions of the xanthophyll cycle pigments violaxanthin and zeaxanthin) were characterized throughout single days and nights to entire seasons in plants growing naturally in contrasting light and temperature environments. All pronounced decreases of intrinsic PSII efficiency took place in the presence of zeaxanthin. The reversibility of these PSII efficiency changes varied widely, ranging from reversible-within-seconds (in a vine experiencing multiple sunflecks under a eucalypt canopy) to apparently permanently locked-in for entire seasons (throughout the whole winter in a subalpine conifer forest at 3,000 m). While close association between low intrinsic PSII efficiency and zeaxanthin accumulation was ubiquitous, accompanying features (such as trans-thylakoid pH gradient, thylakoid protein composition, and phosphorylation) differed among contrasting conditions. The strongest and longest-lasting depressions in intrinsic PSII efficiency were seen in the most stress-tolerant species. Evergreens, in particular, showed the most pronounced modulation of PSII efficiency and thermal dissipation, and are therefore suggested as model species for the study of photoprotection. Implications of the responses of field-grown plants in nature for mechanistic models are discussed.

Photosynthetic Quantum Yield Dynamics: From Photosystems to Leaves

Abstract: The mechanisms underlying the wavelength dependence of the quantum yield for CO 2 fixation (a) and its acclimation to the growth-light spectrum are quantitatively addressed, combining in vivo physiological and in vitro molecular methods. Cucumber (Cucumis sativus) was grown under an artificial sunlight spectrum, shade light spectrum, and blue light, and the quantum yield for photosystem I (PSI) and photosystem II (PSII) electron transport and a were simultaneously measured in vivo at 20 different wavelengths. The wavelength dependence of the photosystem excitation balance was calculated from both these in vivo data and in vitro from the photosystem composition and spectroscopic properties. Measuring wavelengths overexciting PSI produced a higher a for leaves grown under the shade light spectrum (i.e., PSI light), whereas wavelengths overexciting PSII produced a higher a for the sun and blue leaves. The shade spectrum produced the lowest PSI:PSII ratio. The photosystem excitation balance calculated from both in vivo and in vitro data was substantially similar and was shown to determine a at those wavelengths where absorption by carotenoids and nonphotosynthetic pigments is insignificant (i.e., >580 nm). We show quantitatively that leaves acclimate their photosystem composition to their growth light spectrum and how this changes the wavelength dependence of the photosystem excitation balance and quantum yield for CO2 fixation. This also proves that combining different wavelengths can enhance quantum yields substantially.

Detailing the optimality of photosynthesis in cyanobacteria through systems biology analysis

Abstract: Photosynthesis has recently gained considerable attention for its potential role in the development of renewable energy sources. Optimizing photosynthetic organisms for biomass or biofuel production will therefore require a systems understanding of photosynthetic processes. We reconstructed a high-quality genome-scale metabolic network for Synechocystis sp. PCC6803 that describes key photosynthetic processes in mechanistic detail. We performed an exhaustive in silico analysis of the reconstructed photosynthetic process under different light and inorganic carbon (Ci) conditions as well as under genetic perturbations. Our key results include the following. (i) We identified two main states of the photosynthetic apparatus: a Ci-limited state and a light-limited state. (ii) We discovered nine alternative electron flow pathways that assist the photosynthetic linear electron flow in optimizing the photosynthesis performance. (iii) A high degree of cooperativity between alternative pathways was found to be critical for optimal autotrophic metabolism. Although pathways with high photosynthetic yield exist for optimizing growth under suboptimal light conditions, pathways with low photosynthetic yield guarantee optimal growth under excessive light or Ci limitation. (iv) Photorespiration was found to be essential for the optimal photosynthetic process, clarifying its role in high-light acclimation. Finally, (v) an extremely high photosynthetic robustness drives the optimal autotrophic metabolism at the expense of metabolic versatility and robustness. The results and modeling approach presented here may promote a better understanding of the photosynthetic process. They can also guide bioengineering projects toward optimal biofuel production in photosynthetic organisms.

Algal evolution in relation to atmospheric CO2: carboxylases, carbon-concentrating mechanisms and carbon oxidation cycles

Abstract: Oxygenic photosynthesis evolved at least 2.4 Ga; all oxygenic organisms use the ribulose bisphosphate carboxylase-oxygenase (Rubisco)–photosynthetic carbon reduction cycle (PCRC) rather than one of the five other known pathways of autotrophic CO2 assimilation. The high CO2 and (initially) O2-free conditions permitted the use of a Rubisco with a high maximum specific reaction rate. As CO2 decreased and O2 increased, Rubisco oxygenase activity increased and 2-phosphoglycolate was produced, with the evolution of pathways recycling this inhibitory product to sugar phosphates. Changed atmospheric composition also selected for Rubiscos with higher CO2 affinity and CO2/O2 selectivity correlated with decreased CO2-saturated catalytic capacity and/or for CO2-concentrating mechanisms (CCMs). These changes increase the energy, nitrogen, phosphorus, iron, zinc and manganese cost of producing and operating Rubisco–PCRC, while biosphere oxygenation decreased the availability of nitrogen, phosphorus and iron. The majority of algae today have CCMs; the timing of their origins is unclear. If CCMs evolved in a low-CO2 episode followed by one or more lengthy high-CO2 episodes, CCM retention could involve a combination of environmental factors known to favour CCM retention in extant organisms that also occur in a warmer high-CO2 ocean. More investigations, including studies of genetic adaptation, are needed.

Silicon nutrition increases grain yield, which, in turn, exerts a feed-forward stimulation of photosynthetic rates via enhanced mesophyll conductance and alters primary metabolism in rice.

Abstract: Silicon (Si) is not considered to be an essential element for higher plants and is believed to have no effect on primary metabolism in unstressed plants. In rice (Oryza sativa), Si nutrition improves grain production; however, no attempt has been made to elucidate the physiological mechanisms underlying such responses. Here, we assessed crop yield and combined advanced gas exchange analysis with carbon isotope labelling and metabolic profiling to measure the effects of Si nutrition on rice photosynthesis, together with the associated metabolic changes, by comparing wild-type rice with the low-Si rice mutant lsi1 under unstressed conditions. Si improved the harvest index, paralleling an increase in nitrogen use efficiency. Higher crop yields associated with Si nutrition exerted a feed-forward effect on photosynthesis which was fundamentally associated with increased mesophyll conductance. By contrast, Si nutrition did not affect photosynthetic gas exchange during the vegetative growth phase or in de-grained plants. In addition, Si nutrition altered primary metabolism by stimulating amino acid remobilization. Our results indicate a stimulation of the source capacity, coupled with increased sink demand, in Si-treated plants; therefore, we identify Si nutrition as an important target in attempts to improve the agronomic yield of rice.

Rubisco activase is a key regulator of non-steady-state photosynthesis at any leaf temperature and, to a lesser extent, of steady-state photosynthesis at high temperature

Abstract: The role of Rubisco activase in steady-state and non-steady-state photosynthesis was analyzed in wild-type (Oryza sativa) and transgenic rice that expressed different amounts of Rubisco activase. Below 25°C, the Rubisco activation state and steady-state photosynthesis were only affected when Rubisco activase was reduced by more than 70%. However, at 40°C, smaller reductions in Rubisco activase content were linked to a reduced Rubisco activation state and steady-state photosynthesis. As a result, overexpression of maize Rubisco activase in rice did not lead to an increase of the Rubisco activation state, nor to an increase in photosynthetic rate below 25°C, but had a small stimulatory effect at 40°C. On the other hand, the rate at which photosynthesis approached the steady state following an increase in light intensity was rapid in Rubisco activase-overexpressing plants, intermediate in the wild-type, and slowest in antisense plants at any leaf temperature. In Rubisco activase-overexpressing plants, Rubisco activation state at low light was maintained at higher levels than in the wild-type. Thus, rapid regulation by Rubisco activase following an increase in light intensity and/or maintenance of a high Rubisco activation state at low light would result in a rapid increase in Rubisco activation state and photosynthetic rate following an increase in light intensity. It is concluded that Rubisco activase plays an important role in the regulation of non-steady-state photosynthesis at any leaf temperature and, to a lesser extent, of steady-state photosynthesis at high temperature.

Optimization of photosynthetic light energy utilization by microalgae

Abstract: Over 50% of the energy losses associated with the conversion of solar energy into chemical energy during photosynthesis are attributed to kinetic constraints between the fast rate of photon capture by the light harvesting apparatus and the slower downstream rate of photosynthetic electron transfer. At full sunlight intensities, energy flux from the light harvesting antennae to the reaction centers may be 100-folds greater than the overall linear electron flow resulting in the dissipation of up to 75% of the captured energy as heat or fluorescence. One possible means to couple energy capture and photosynthetic electron transfer more efficiently is to reduce the optical cross-section of the light harvesting antennae. We show that by partially reducing chlorophyll b levels in the green alga, Chlamydomonas reinhardtii, we can tune the peripheral light harvesting antennae size for increased photosynthetic efficiency resulting in more than a two-fold increase in photosynthetic rate at high light intensities and a 30% increase in growth rate at saturating light intensities. Unlike chlorophyll b-less mutants which lack the peripheral light harvesting antennae; transgenics with intermediate sized peripheral antennae have the advantage that they can carry out state transitions facilitating enhanced cyclic ATP synthesis and have robust zeaxanthin–violaxanthin cycles providing protection from high light levels. It is hypothesized that the large antennae size of wild-type algae and land plants offers a competitive advantage in mixed cultures due to the ability of photosynthetic organisms with large light harvesting antennae to shade competing species and to harvest light at low flux densities.

Modelling photosynthetic responses to temperature of grapevine (Vitis vinifera cv. Semillon) leaves on vines grown in a hot climate.

Abstract: Field measurements of photosynthesis of Vitis vinifera cv. Semillon leaves in relation to a hot climate, and responses to photon flux densities (PFDs) and internal CO(2) concentrations (c(i) ) at leaf temperatures from 20 to 40 °C were undertaken. Average rates of photosynthesis measured in situ decreased with increasing temperature and were 60% inhibited at 45 °C compared with 25 °C. This reduction in photosynthesis was attributed to 15-30% stomatal closure. Light response curves at different temperatures revealed light-saturated photosynthesis optimal at 30 °C but also PFDs saturating photosynthesis increased from 550 to 1200 µmol (photons) m(-2)s(-1) as temperatures increased. Photosynthesis under saturating CO(2) concentrations was optimal at 36 °C while maximum rates of ribulose 1,5-bisphosphate (RuBP) carboxylation (V(cmax)) and potential maximum electron transport rates (J(max)) were also optimal at 39 and 36 °C, respectively. Furthermore, the high temperature-induced reduction in photosynthesis at ambient CO(2) was largely eliminated. The chloroplast CO(2) concentration at the transition from RuBP regeneration to RuBP carboxylation-limited assimilation increased steeply with an increase in leaf temperature. Semillon assimilation in situ was limited by RuBP regeneration below 30 °C and above limited by RuBP carboxylation, suggesting high temperatures are detrimental to carbon fixation in this species.

Proteaceae from severely phosphorus-impoverished soils extensively replace phospholipids with galactolipids and sulfolipids during leaf development to achieve a high photosynthetic phosphorus-use-efficiency.

Abstract: Summary Proteaceae species in south-western Australia occur on severely phosphorus (P)-impoverished soils. They have very low leaf P concentrations, but relatively fast rates of photosynthesis, thus exhibiting extremely high photosynthetic phosphorus-use-efficiency (PPUE). Although the mechanisms underpinning their high PPUE remain unknown, one possibility is that these species may be able to replace phospholipids with nonphospholipids during leaf development, without compromising photosynthesis. For six Proteaceae species, we measured soil and leaf P concentrations and rates of photosynthesis of both young expanding and mature leaves. We also assessed the investment in galactolipids, sulfolipids and phospholipids in young and mature leaves, and compared these results with those on Arabidopsis thaliana, grown under both P-sufficient and P-deficient conditions. In all Proteaceae species, phospholipid levels strongly decreased during leaf development, whereas those of galactolipids and sulfolipids strongly increased. Photosynthetic rates increased from young to mature leaves. This shows that these species extensively replace phospholipids with nonphospholipids during leaf development, without compromising photosynthesis. A considerably less pronounced shift was observed in A. thaliana. Our results clearly show that a low investment in phospholipids, relative to nonphospholipids, offers a partial explanation for a high photosynthetic rate per unit leaf P in Proteaceae adapted to P-impoverished soils.

Increased fructose 1,6-bisphosphate aldolase in plastids enhances growth and photosynthesis of tobacco plants

Abstract: The Calvin cycle is the initial pathway of photosynthetic carbon fixation, and several of its reaction steps are suggested to exert rate-limiting influence on the growth of higher plants. Plastid fructose 1,6-bisphosphate aldolase (aldolase, EC 4.1.2.13) is one of the nonregulated enzymes comprising the Calvin cycle and is predicted to have the potential to control photosynthetic carbon flux through the cycle. In order to investigate the effect of overexpression of aldolase, this study generated transgenic tobacco (Nicotiana tabacum L. cv Xanthi) expressing Arabidopsis plastid aldolase. Resultant transgenic plants with 1.4-1.9-fold higher aldolase activities than those of wild-type plants showed enhanced growth, culminating in increased biomass, particularly under high CO₂ concentration (700 ppm) where the increase reached 2.2-fold relative to wild-type plants. This increase was associated with a 1.5-fold elevation of photosynthetic CO₂ fixation in the transgenic plants. The increased plastid aldolase resulted in a decrease in 3-phosphoglycerate and an increase in ribulose 1,5-bisphosphate and its immediate precursors in the Calvin cycle, but no significant changes in the activities of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) or other major enzymes of carbon assimilation. Taken together, these results suggest that aldolase overexpression stimulates ribulose 1,5-bisphosphate regeneration and promotes CO₂ fixation. It was concluded that increased photosynthetic rate was responsible for enhanced growth and biomass yields of aldolase-overexpressing plants.

Heterosis of Arabidopsis hybrids between C24 and Col is associated with increased photosynthesis capacity

Abstract: Arabidopsis thaliana shows hybrid vigor (heterosis) in progeny of crosses between Columbia-0 and C24 accessions. Hybrid vigor was evident as early as mature seeds and in seedlings 3 d after sowing (DAS). At 3 DAS, genes encoding chloroplast-located proteins were significantly overrepresented (187) among the 724 genes that have greater than midparent values of expression in the hybrid. Many of these genes are involved in chlorophyll biosynthesis and photosynthesis. The rate of photosynthesis was constant per unit leaf area in parents and hybrids. Larger cell sizes in the hybrids were associated with more chloroplasts per cell, more total chlorophyll, and more photosynthesis. The increased transcription of the chloroplast-targeted genes was restricted to the 3–7 DAS period. At 10 DAS, only 118 genes had expression levels different from the expected midparent value in the hybrid, and only 12 of these genes were differentially expressed at 3 DAS. The early increase in activity of genes involved in photosynthesis and the associated phenomena of increases in cell size and number through development, leading to larger leaf areas of all leaves in the hybrid, suggest a central role for increased photosynthesis in the production of the heterotic biomass. In support of this correlation, we found that an inhibitor of photosynthesis eliminated heterosis and that higher light intensities enhanced both photosynthesis and heterosis. In hybrids with low-level heterosis (Landsberg erecta x Columbia-0), chloroplast-targeted genes were not up-regulated and leaf areas were only marginally increased.

Co-ordination of hydraulic and stomatal conductances across light qualities in cucumber leaves

Abstract: Long-term effects of light quality on leaf hydraulic conductance (Kleaf) and stomatal conductance (gs) were studied in cucumber, and their joint impact on leaf photosynthesis in response to osmotic-induced water stress was assessed. Plants were grown under low intensity monochromatic red (R, 640 nm), blue (B, 420 nm) or combined red and blue (R:B, 70:30) light. Kleaf and gs were much lower in leaves that developed without blue light. Differences in gs were caused by differences in stomatal aperture and stomatal density, of which the latter was largely due to differences in epidermal cell size and hardly due to stomatal development. Net photosynthesis (AN) was lowest in R-, intermediate in B-, and highest in RB- grown leaves. The low AN in R-grown leaves correlated with a low leaf internal CO2 concentration and reduced PSII operating efficiency. In response to osmotic stress, all leaves showed similar degrees of stomatal closure, but the reduction in AN was larger in R- than in B- and RB-grown leaves. This was probably due to damage of the photosynthetic apparatus, which only occurred in R-grown leaves. The present study shows the co-ordination of Kleaf and gs across different light qualities, while the presence of blue in the light spectrum seems to drive both Kleaf and gs towards high, sun-type leaf values, as was previously reported for maximal photosynthetic capacity and leaf morphology. The present results suggest the involvement of blue light receptors in the usually harmonized development of leaf characteristics related to water relations and photosynthesis under different light environments.

Manganese-oxidizing photosynthesis before the rise of cyanobacteria

Abstract: The emergence of oxygen-producing (oxygenic) photosynthesis fundamentally transformed our planet; however, the processes that led to the evolution of biological water splitting have remained largely unknown. To illuminate this history, we examined the behavior of the ancient Mn cycle using newly obtained scientific drill cores through an early Paleoproterozoic succession (2.415 Ga) preserved in South Africa. These strata contain substantial Mn enrichments (up to ∼17 wt %) well before those associated with the rise of oxygen such as the ∼2.2 Ga Kalahari Mn deposit. Using microscale X-ray spectroscopic techniques coupled to optical and electron microscopy and carbon isotope ratios, we demonstrate that the Mn is hosted exclusively in carbonate mineral phases derived from reduction of Mn oxides during diagenesis of primary sediments. Additional observations of independent proxies for O2—multiple S isotopes (measured by isotope-ratio mass spectrometry and secondary ion mass spectrometry) and redox-sensitive detrital grains—reveal that the original Mn-oxide phases were not produced by reactions with O2, which points to a different high-potential oxidant. These results show that the oxidative branch of the Mn cycle predates the rise of oxygen, and provide strong support for the hypothesis that the water-oxidizing complex of photosystem II evolved from a former transitional photosystem capable of single-electron oxidation reactions of Mn.

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

Abstract: 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.

Alternating electron and proton transfer steps in photosynthetic water oxidation

Abstract: Water oxidation by cyanobacteria, algae, and plants is pivotal in oxygenic photosynthesis, the process that powers life on Earth, and is the paradigm for engineering solar fuel–production systems. Each complete reaction cycle of photosynthetic water oxidation requires the removal of four electrons and four protons from the catalytic site, a manganese–calcium complex and its protein environment in photosystem II. In time-resolved photothermal beam deflection experiments, we monitored apparent volume changes of the photosystem II protein associated with charge creation by light-induced electron transfer (contraction) and charge-compensating proton relocation (expansion). Two previously invisible proton removal steps were detected, thereby filling two gaps in the basic reaction-cycle model of photosynthetic water oxidation. In the S2 → S3 transition of the classical S-state cycle, an intermediate is formed by deprotonation clearly before electron transfer to the oxidant (). The rate-determining elementary step (τ, approximately 30 µs at 20 °C) in the long-distance proton relocation toward the protein–water interface is characterized by a high activation energy (Ea = 0.46 ± 0.05 eV) and strong H/D kinetic isotope effect (approximately 6). The characteristics of a proton transfer step during the S0 → S1 transition are similar (τ, approximately 100 µs; Ea = 0.34 ± 0.08 eV; kinetic isotope effect, approximately 3); however, the proton removal from the Mn complex proceeds after electron transfer to . By discovery of the transient formation of two further intermediate states in the reaction cycle of photosynthetic water oxidation, a temporal sequence of strictly alternating removal of electrons and protons from the catalytic site is established.