Characteristics of C-4 photosynthesis in stems and petioles of C-3 flowering plants

Most plants are known as C-3 plants because the first product of photosynthetic CO2 fixation is a three-carbon compound. C-4 plants, which use an alternative pathway in which the first product is a four-carbon compound, have evolved independently many times and are found in at least 18 families. In addition to differences in their biochemistry, photosynthetic organs of C-4 plants show alterations in their anatomy and ultrastructure. Little is known about whether the biochemical or anatomical characteristics of C-4 photosynthesis evolved first. Here we report that tobacco, a typical C-3 plant, shows characteristics of C-4 photosynthesis in cells of stems and petioles that surround the xylem and phloem, and that these cells are supplied with carbon for photosynthesis from the vascular system and not from stomata. These photosynthetic cells possess high activities of enzymes characteristic of C-4 photosynthesis, which allow the decarboxylation of four-carbon organic acids from the xylem and phloem, thus releasing CO2 for photosynthesis. These biochemical characteristics of C-4 photosynthesis in cells around the vascular bundles of stems of C-3 plants might explain why C-4 photosynthesis has evolved independently many times

Spectral signatures of photosynthesis II: coevolution with other stars and the atmosphere on extrasolar worlds

As photosynthesis on Earth produces the primary signatures of life that can be detected astronomically at the global scale, a strong focus of the search for extrasolar life will be photosynthesis, particularly photosynthesis that has evolved with a different parent star. We take planetary atmospheric compositions simulated by Segura, et al. (2003, 2005) for Earth-like planets around observed F2V and K2V stars, modeled M1V and M5V stars, and around the active M4.5V star AD Leo; our scenarios use Earth's atmospheric composition as well as very low O2 content in case anoxygenic photosynthesis dominates. We calculate the incident spectral photon flux densities at the surface of the planet and under water. We identify bands of available photosynthetically relevant radiation and find that photosynthetic pigments on planets around F2V stars may peak in absorbance in the blue, K2V in the red-orange, and M stars in the NIR, in bands at 0.93-1.1 microns, 1.1-1.4 microns, 1.5-1.8 microns, and 1.8-2.5 microns. In addition, we calculate wavelength restrictions for underwater organisms and depths of water at which they would be protected from UV flares in the early life of M stars. We estimate the potential productivity for both surface and underwater photosynthesis, for both oxygenic and anoxygenic photosynthesis, and for hypothetical photosynthesis in which longer wavelength, multi-photosystem series are used.Comment: 59 pages, 4 figures, 4 tables, forthcoming in Astrobiology ~March 200

Phenotypic landscape inference reveals multiple evolutionary paths to C4_4 photosynthesis

C$_4$ photosynthesis has independently evolved from the ancestral C$_3$ 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 C$_4$ 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 C$_3$, C$_4$, or intermediate C$_3$-C$_4$ 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 C$_4$ phenotype evolved. The alternative evolutionary histories underlying the appearance of C$_4$ photosynthesis were determined by ancestral lineage and initial phenotypic alterations unrelated to photosynthesis. We conclude that the order of C$_4$ trait acquisition is flexible and driven by non-photosynthetic drivers. This flexibility will have facilitated the convergent evolution of this complex trait

Does Long-Term Elevation of CO2 Concentration Increase Photosynthesis in Forest Floor Vegetation? (Indiana Strawberry in a Maryland Forest).

As the partial pressure of CO2 (pCO2) in the atmosphere rises, photorespiratory loss of carbon in C3 photosynthesis will diminish and the net efficiency of light-limited photosynthetic carbon uptake should rise. We tested this expectation for Indiana strawberry (Duchesnea indica) growing on a Maryland forest floor. Open-top chambers were used to elevate the pCO2 of a forest floor habitat to 67 Pa and were paired with control chambers providing an ambient pCO2 of 38 Pa. After 3.5 years, D. indica leaves grown and measured in the elevated pCO2 showed a significantly greater maximum quantum efficiency of net photosynthesis (by 22%) and a lower light compensation point (by 42%) than leaves grown and measured in the control chambers. The quantum efficiency to minimize photorespiration, measured in 1% O2, was the same for controls and plants grown at elevated pCO2. This showed that the maximum efficiency of light-energy transduction into assimilated carbon was not altered by acclimation and that the increase in light-limited photosynthesis at elevated pCO2 was simply a function of the decrease in photorespiration. Acclimation did decrease the ribulose-1,5-bisphosphate carboxylase/oxygenase and light-harvesting chlorophyll protein content of the leaf by more than 30%. These changes were associated with a decreased capacity for light-saturated, but not light-limited, photosynthesis. Even so, leaves of D. indica grown and measured at elevated pCO2 showed greater light-saturated photosynthetic rates than leaves grown and measured at the current atmospheric pCO2. In situ measurements under natural forest floor lighting showed large increases in leaf photosynthesis at elevated pCO2, relative to controls, in both summer and fall. The increase in efficiency of light-limited photosynthesis with elevated pCO2 allowed positive net photosynthetic carbon uptake on days and at locations on the forest floor that light fluxes were insufficient for positive net photosynthesis in the current atmospheric pCO2

Photosynthetic responses of three common mosses from continental Antarctica

Predicting the effects of climate change on Antarctic terrestrial vegetation requires a better knowledge of the ecophysiology of common moss species. In this paper we provide a comprehensive matrix for photosynthesis and major environmental parameters for three dominant Antarctic moss species (Bryum subrotundifolium, B. pseudotriquetrum and Ceratodon purpureus). Using locations in southern Victoria Land, (Granite Harbour, 77°S) and northern Victoria Land (Cape Hallett, 72°S) we determined the responses of net photosynthesis and dark respiration to thallus water content, thallus temperature, photosynthetic photon flux densities and CO2 concentration over several summer seasons. The studies also included microclimate recordings at all sites where the research was carried out in field laboratories. Plant temperature was influenced predominantly by the water regime at the site with dry mosses being warmer. Optimal temperatures for net photosynthesis were 13.7°C, 12.0°C and 6.6°C for B. subrotundifolium, B. pseudotriquetrum and C. purpureus, respectively and fall within the known range for Antarctic mosses. Maximal net photosynthesis at 10°C ranked as B. subrotundifolium > B. pseudotriquetrum > C. purpureus. Net photosynthesis was strongly depressed at subzero temperatures but was substantial at 0°C. Net photosynthesis of the mosses was not saturated by light at optimal water content and thallus temperature. Response of net photosynthesis to increase in water content was as expected for mosses although B. subrotundifolium showed a large depression (60%) at the highest hydrations. Net photosynthesis of both B. subrotundifolium and B. pseudotriquetrum showed a large response to increase in CO2 concentration and this rose with increase in temperature; saturation was not reached for B. pseudotriquetrum at 20°C. There was a high level of variability for species at the same sites in different years and between different locations. This was substantial enough to make prediction of the effects of climate change very difficult at the moment

Understanding plant responses to drought: how important is woody tissue photosynthesis?

Within trees, it is known that a part of the respired CO2 is assimilated in chlorophyll-containing stem and branch tissues. However, the role of this woody tissue photosynthesis in tree functioning remains unclear, in particular under drought stress conditions. In this study, stem diameter and leaf photosynthesis were measured for one-year-old cutting-derived plants of Populus nigra 'Monviso' under both well-watered and drought stress conditions. Half of the plants were subjected to a stem and branch light-exclusion treatment to prevent woody tissue photosynthesis to occur, while the other trees served as controls. Drought stress was induced in both treatments by limiting the water supply. We found that under well-watered conditions, light-exclusion resulted in reduced stem radial daily growth rate (DG) relative to DG observed for control trees. In response to drought, stem shrinkage of the light-excluded trees was more pronounced as compared to the control trees. Maximum leaf net photosynthesis (A(max)) decreased more rapidly in light-excluded trees compared to the controls during drought stress. Our results are the first to report on the potentially significant role of woody tissue photosynthesis in tree drought stress tolerance. Moreover, our study implies that the impact of assimilation of respired CO2 on tree functioning extends beyond local stem processes and indicates that woody tissue photosynthesis is potentially a key factor in understanding plant responses to drought stress

Development of a portable leaf photosynthesis and volatile organic compounds emission system.

Understanding how plant carbon metabolism responds to environmental variables such as light is central to understanding ecosystem carbon cycling and the production of food, biofuels, and biomaterials. Here, we couple a portable leaf photosynthesis system to an autosampler for volatile organic compounds (VOCs) to enable field observations of net photosynthesis simultaneously with emissions of VOCs as a function of light. Following sample collection, VOCs are analyzed using automated thermal desorption-gas chromatograph-mass spectrometry (TD-GC-MS). An example is presented from a banana plant in the central Amazon with a focus on the response of photosynthesis and the emissions of eight individual monoterpenes to light intensity. Our observations reveal that banana leaf emissions represent a 1.1 +/- 0.1% loss of photosynthesis by carbon. Monoterpene emissions from banana are dominated by trans-β-ocimene, which accounts for up to 57% of total monoterpene emissions at high light. We conclude that the developed system is ideal for the identification and quantification of VOC emissions from leaves in parallel with CO2 and water fluxes.The system therefore permits the analysis of biological and environmental sensitivities of carbon metabolism in leaves in remote field locations, resulting in the emission of hydrocarbons to the atmosphere.•A field-portable system is developed for the identification and quantification of VOCs from leaves in parallel with leaf physiological measurements including photosynthesis and transpiration.•The system will enable the characterization of carbon and energy allocation to the biosynthesis and emission of VOCs linked with photosynthesis (e.g. isoprene and monoterpenes) and their biological and environmental sensitivities (e.g. light, temperature, CO2).•Allow the development of more accurate mechanistic global VOC emission models linked with photosynthesis, improving our ability to predict how forests will respond to climate change. It is our hope that the presented system will contribute with critical data towards these goals across Earth's diverse tropical forests

Ecophysiological traits of grasses: resolving the effects of photosynthetic pathway and phylogeny

C4 photosynthesis is an important example of convergent evolution in plants, having arisen in eudicots, monocots and diatoms. Comparisons between such diverse groups are confounded by phylogenetic and ecological differences, so that only broad generalisations can be made about the role of C4 photosynthesis in
determining ecophysiological traits. However, 60% of C4 species occur in the grasses (Poaceae) and molecular phylogenetic techniques confirm that there are between 8 and 17 independent origins of C4 photosynthesis in the Poaceae. In a screening experiment, we compared leaf physiology and growth traits across several major
independent C3 & C4 groups within the Poaceae, asking 1) which traits differ consistently between photosynthetic
types and 2) which traits differ consistently between clades within each photosynthetic type

Evolutionary temperature compensation of carbon fixation in marine phytoplankton

The efficiency of carbon sequestration by the biological pump could decline in the coming decades because respiration tends to increase more with temperature than photosynthesis. Despite these differences in the short-term temperature sensitivities of photosynthesis and respiration, it remains unknown whether the long-term impacts of global warming on metabolic rates of phytoplankton can be modulated by evolutionary adaptation. We found that respiration was consistently more temperature dependent than photosynthesis across 18 diverse marine phytoplankton, resulting in universal declines in the rate of carbon fixation with short-term increases in temperature. Long-term experimental evolution under high temperature reversed the short-term stimulation of metabolic rates, resulting in increased rates of carbon fixation. Our findings suggest that thermal adaptation may therefore have an ameliorating impact on the efficiency of phytoplankton as primary mediators of the biological carbon pump