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DOE OSTI2020

Triose phosphate utilization limitation: an unnecessary complexity in terrestrial biosphere model representation of photosynthesis

Triose phosphates are the principal product of photosynthesis. They are used within the chloroplast for starch synthesis, or translocated to the cytosol where they are used to fuel sucrose synthesis. Use of triose phosphate releases inorganic phosphate, and is under strict metabolic control that matches the supply of triose phosphate from the Calvin-Benson cycle to the demand for carbon by sinks (Heldt & Heldt, 2011; McClain & Sharkey, 2019). However, a low rate of triose phosphate utilization (TPU) can deplete the phosphate pool, restrict ATP synthesis and reduce the availability of ATP to power the Calvin-Benson cycle, thereby limiting photosynthesis (Sharkey, 1985).

54 ENVIRONMENTAL SCIENCES↗

DOE OSTI2026

Disruption of starch biosynthesis and triacylglycerol degradation impairs growth but improves photosynthesis in Arabidopsis

Enhancing lipid accumulation by redirecting carbon from starch to triacylglycerol (TAG) in vegetative tissues is a promising strategy for developing high-energy-density crops for bioenergy production. However, our understanding of how starch and TAG metabolism interact and how this interaction affects growth and photosynthesis is incomplete. Here, we investigated the metabolic and physiological consequences of disrupting starch biosynthesis and TAG turnover in Arabidopsis thaliana by generating single, double, and triple mutants involving ADG1 (starch biosynthesis), TGD1 (lipid trafficking), and SDP1\\\\\\\\r\\\\\\\\n(TAG lipase). Unexpectedly, elimination of starch biosynthesis lowered TAG levels in the high-TAG-accumulating tgd1 mutant, primarily through enhanced TAG breakdown. This decline was completely reversed by sucrose supplementation, suggesting that TAG degradation was induced by carbon limitation.\\\\\\\\r\\\\\\\\nGenetic suppression of TAG turnover via SDP1 disruption in the starchless tgd1 adg1-1 background led to a nine-fold rise in leaf TAG accumulation in the tgd1 sdp1 adg1-1 triple mutant, together with enhanced photosynthetic performance. However, this metabolic reprogramming incurred growth penalties.\\\\\\\\r\\\\\\\\nOur results highlight the role of dynamic TAG turnover in maintaining metabolic balance and photosynthesis in starch-deficient backgrounds. These findings underscore the need for refined metabolic engineering strategies that coordinate TAG biosynthesis and degradation to optimize lipid accumulation in bioenergy crops.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2022

Stomatal conductance, not biochemistry, drives low temperature acclimation of photosynthesis in Populus balsamifera , regardless of nitrogen availability

Low temperature thermal acclimation may require adjustments to nitrogen and water use to sustain photosynthesis due to slow enzyme functioning and high-water viscosity. However, understanding of photosynthetic acclimation to temperatures below 11°C is limited. Here, we acclimated Populus balsamifera to 6 °C and 10 °C (6A and 10A, respectively) and provided the trees with either high or low N fertilizer. We measured net CO 2 assimilation rates (A net ), stomatal conductance (g s ), maximum rates of Rubisco carboxylation (V cmax ) and electron transport (J max ), and dark respiration (R d ) at leaf temperatures of 2, 6, 10, 14 and 18 °C, along with leaf N concentrations. The 10A trees had higher A net than the 6A trees at warmer leaf temperatures, which was correlated with higher g s in the 10A trees. The instantaneous temperature responses of V cmax , J max and R d were similar for trees from both acclimation temperatures. While soil N availability increased leaf N concentrations, this had no effect on acclimation of photosynthesis or respiration. Our results indicate that acclimation below 11°C occurred primarily through changes in stomatal conductance, not photosynthetic biochemistry, and was unaffected by short-term N supply. Thermal acclimation of stomatal conductance should therefore be a priority for future carbon cycle model development.

54 ENVIRONMENTAL SCIENCES↗

DOE OSTI2020

The structure of Photosystem I acclimated to far-red light illuminates an ecologically important acclimation process in photosynthesis

Phototrophic organisms are superbly adapted to different light environments but often must acclimate to challenging competition for visible light wavelengths in their niches. Some cyanobacteria overcome this challenge by expressing paralogous photosynthetic proteins and by synthesizing and incorporating ~8% chlorophyll f into their Photosystem I (PSI) complexes, enabling them to grow under far-red light (FRL). We solved the structure of FRL-acclimated PSI from the cyanobacterium Fischerella thermalis PCC 7521 by single-particle, cryo–electron microscopy to understand its structural and functional differences. Four binding sites occupied by chlorophyll f are proposed. Subtle structural changes enable FRL-adapted PSI to extend light utilization for oxygenic photosynthesis to nearly 800 nm. This structure provides a platform for understanding FRL-driven photosynthesis and illustrates the robustness of adaptive and acclimation mechanisms in nature.

Science & Technology - Other Topics↗

DOE OSTI2021

Isotope ratio-based quantification of carbon assimilation highlights the role of plastidial isoprenoid precursor availability in photosynthesis

Abstract Background We report a method to estimate carbon assimilation based on isotope ratio-mass spectrometry (IRMS) of 13 CO 2 labeled plant tissue. Photosynthetic carbon assimilation is the principal experimental observable which integrates important aspects of primary plant metabolism. It is traditionally measured through gas exchange. Despite its centrality in plant research, gas exchange performs poorly with rosette growth habits typical of Arabidopsis thaliana , mutant lines with limited biomass, and accounts poorly for leaf shading. Results IRMS-based carbon assimilation values from plants labeled at different light intensities were compared to those obtained by gas exchange, and the two methods yielded similar values. Using this method, we observed a strong correlation between 13 C content and labeling time (R 2 = 0.999) for 158 wild-type plants labeled for 6 to 42 min. Plants cultivated under different light regimes showed a linear response with respect to carbon assimilation, varying from 7.38 nmol 13 C mg −1 leaf tissue min −1 at 80 PAR to 19.27 nmol 13 C mg −1 leaf tissue min −1 at 500 PAR. We applied this method to examine the link between inhibition of the 2 C -methyl- d -erythritol-4-phosphate (MEP) pathway and suppression of photosynthesis. A significant decrease in carbon assimilation was observed when metabolic activity in the MEP pathway was compromised by mutation or herbicides targeting the MEP pathway. Mutants affected in MEP pathway genes 1-DEOXY- d -XYLULOSE 5-PHOSPHATE SYNTHASE (DXS) or 1-HYDROXY-2-METHYL-2-(E)-BUTENYL 4-DIPHOSPHATE SYNTHASE (HDS) showed assimilation rates 36% and 61% lower than wild type. Similarly, wild type plants treated with the MEP pathway inhibitors clomazone or fosmidomycin showed reductions of 52% and 43%, respectively, while inhibition of the analogous mevalonic acid pathway, which supplies the same isoprenoid intermediates in the cytosol, did not, suggesting inhibition of photosynthesis was specific to disruption of the MEP pathway. Conclusions This method provides an alternative to gas exchange that offers several advantages: resilience to differences in leaf overlap, measurements based on tissue mass rather than leaf surface area, and compatibility with mutant Arabidopsis lines which are not amenable to gas exchange measurements due to low biomass and limited leaf surface area. It is suitable for screening large numbers of replicates simultaneously as well as post-hoc analysis of previously labeled plant tissue and is complementary to downstream detection of isotopic label in targeted metabolite pools.

2C‑methyl‑d‑erythritol 4‑phosphate pathway↗

DOE OSTI2024

Contributions of David Mauzerall to photosynthesis research - celebrating his 95 th birthday

We honor here Professor David Mauzerall, a pioneer in the fields of photochemistry and photobiology of porphyrins and chlorophylls in vitro and in vivo, on the occasion of his 95th birthday. Throughout his career at The Rockefeller University, he refined our understanding of how chlorophyll converts light energy into chemical energy. He exploited top-of-the-line laser technology in developing photoacoustics and a variety of other innovative experimental approaches. His experimental work and conceptual insights contributed greatly to our understanding of photosynthesis and the possible role of photosynthesis in the origin of life. His contributions include many landmark single-authored and collaborative papers, and his legacy includes the training of others who have become authorities themselves. After providing a brief description of his research accomplishments, we include tributes from several of his coworkers and his daughters highlighting their valuable experiences with David Mauzerall on this milestone birthday.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2022

Exogenous Hemin alleviated cadmium stress in maize (Zea mays L.) by enhancing leaf photosynthesis, AsA-GSH cycle and polyamine metabolism

Cadmium (Cd) stress is one of the principal abiotic stresses that inhibit maize growth. The research was to explore (hemin chloride) Hemin (100 μmol L −1 ) on photosynthesis, ascorbic acid (AsA)-glutathione (GSH) cycle system, and polyamine metabolism of maize under Cd stress (85 mg L −1 ) using nutrient solution hydroponics, with Tiannong 9 (Cd tolerant) and Fenghe 6 (Cd sensitive) as experimental materials. The results showed that Hemin can increase leaf photosynthetic pigment content and ameliorate the ratio of Chlorophyll a/chlorophyll b ( Chla / Chlb ) under Cd stress. The values of ribose 1, 5-diphosphate carboxylase/oxygenase (RuBPcase) and phosphoenolpyruvate carboxylase (PEPCase), and total xanthophyll cycle pool [(violoxanthin (V), antiflavin (A) and zeaxanthin (Z)] increased, which enhancing xanthophyll cycle (DEPS) de-epoxidation, and alleviating stomatal and non-stomatal limitation of leaf photosynthesis. Hemin significantly increased net photosynthetic rate ( P n ), stomatal conductance ( g s ), transpiration rate ( T r ), photochemical quenching coefficient ( qP ), PSII maximum photochemical efficiency ( F v /F m ), and electron transfer rate ( ETR ), which contributed to the improvement of the PSII photosynthetic system. Compared with Cd stress, Hemin can reduce thiobartolic acid reactant (TBARS) content, superoxide anion radical (O 2 − ) production rate, hydrogen peroxide (H 2 O 2 ) accumulation, and the extent of electrolyte leakage (EL); decreased the level of malondialdehyde (MDA) content and increased the activities of superoxide dismutase (SOD), peroxidase (POD) and catalase (CAT); slowed the decrease in dehydroascorbic acid reductase (DHAR) and monodehydroascorbate reductase (MDHAR) activity and the increase in glutathione reductase (GR) and ascorbate peroxidase (APX) activity in leaves; promoted the increase in AsA and GSH content, decreased dehydroascorbic acid (DHA) and oxidized glutathione (GSSG), and increased AsA/DHA and GSH/GSSG ratios under Cd stress. Hemin promoted the increase of conjugated and bound polyamine content, and the conversion process speed of free putrescine (Put) to free spermine (Spm) and spermidine (Spd) in maize; decreased polyamine oxidase (PAO) activity and increased diamine oxidase (DAO), arginine decarboxylase (ADC), ornithine decarboxylase (ODC) and S-adenosylmethionine decarboxylase (SAMDC) enzyme activities in leaves under Cd stress.

Piao, Lin↗

NASA NTRS1988

Relating seasonal patterns of the AVHRR vegetation index to simulated photosynthesis and transpiration of forests in different climates

Weekly AVHRR Normalized Difference Vegetation Index (NDVI) values for 1983-1984 for seven sites of diverse climate in North America were correlated with results of an ecosystem simulation model of a hypothetical forest stand for the corresponding period at each site. The tendency of raw NDVI data to overpredict photosynthesis and transpiration on water limited sites was shown to be partially corrected by using an aridity index of annual radiation/annual precipitation. The results suggest that estimates of vegetation productivity using the global vegetation index are only accurate as annual integrations, unless unsubsampled local area coverage NDVI data can be tested against forest photosynthesis, transpiration and aboveground net primary production data measured at shorter time intervals.

Running, Steven W.↗

NASA NTRS1990

Calculating Irradiance For Photosynthesis In The Ocean

Mathematical model predicts available and usable irradiances. Yields estimates of irradiance available for photosynthesis (Epar) and irradiance usable for photosynthesis (Epur) as functions of depth in ocean. Describes Epur and Epar in terms of spectral parameters measured remotely (from satellites or airplanes). These irradiances useful in studies of photosynthetic productivity of phytoplankton in euphotic layer.

Collins, Donald J.↗

NASA NTRS1992

Nutrient control of phytoplankton photosynthesis in the western North Atlantic

Results from several years of oceanographic cruises are reported which show that the parameters of the photosynthesis-light curve of the flora of the North Sargasso Sea are remarkably constant in magnitude, except during the spring phytoplankton bloom when their magnitudes are noticeably higher. These results are interpreted as providing direct evidence for nutrient control of photosynthesis in the open ocean. The findings also reinforce the plausibility of using biogeochemical provinces to partition the ocean into manageable units for basin- or global-scale analysis. They show that seasonal changes in critical parameter should not be overlooked if robust carbon budgets are to be constructed, and illustrate the value of attacking the parameters that control the key fluxes, rather than the fluxes themselves, when investigating the ocean carbon cycle.

Platt, Trevor↗

NASA NTRS2002

An Optical Index of Phytoplankton Photoacclimation and Its Relation to Light-Saturated Photosynthesis in the Sea

In relation to understanding ocean biology at the global scale, one of NASA's primary foci has been measurements of near-surface concentrations of phytoplankton chlorophyll. Chlorophyll is an important light-absorbing pigment in phytoplankton. The absorbed light energy is used to fix carbon in the process of photosynthesis. Photosynthesis, in turn, is critical to the growth of phytoplankton and the function of entire marine ecosystems. Thus, the use of satellite surface chlorophyll data to estimate primary production in the ocean has been a key focus of much biological oceanography research. One of the major challenges in this research is to develop relationships that allow a given chlorophyll concentration (a standing stock) to be interpreted in terms of carbon fixation (a rate). This problem centers on the description of the light-saturated photosynthetic rate, Pbmax. In this paper, we describe how optical measurements of light attenuation provide information on particulate organic carbon (POC) concentrations. We then show how the ratio of POC to chlorophyll (Theta) provides critical information on variability in Pbmax. We then test this relationship between Theta and Pbmax using field data from a variety of open ocean ecosystems.

Behrenfeld, Michael J.↗

NASA NTRS2002

Growth and photosynthesis of Japanese flowering cherry under simulated microgravity conditions

The photosynthetic rate, the leaf characteristics related to photosynthesis, such as the chlorophyll content, chlorophyll a/b ratio and density of the stomata, the leaf area and the dry weight in seedlings of Japanese flowering cherry grown under normal gravity and simulated microgravity conditions were examined. No significant differences were found in the photosynthetic rates between the two conditions. Moreover, leaf characteristics such as the chlorophyll content, chlorophyll a/b ratio and density of the stomata in the seedlings grown under the simulated microgravity condition were not affected. However, the photosynthetic product of the whole seedling under the simulated microgravity condition increased compared with the control due to its leaf area increase. The results suggest that dynamic gravitational stimulus controls the partitioning of the products of photosynthesis.

Weightlessness Simulation↗

NASA NTRS1999

The effects of UV radiation A and B on diurnal variation in photosynthesis in three taxonomically and ecologically diverse microbial mats

Photosynthetic primary production, the basis of most global food chains, is inhibited by UV radiation. Evaluating UV inhibition is therefore important for assessing the role of natural levels of UV radiation in regulating ecosystem behavior as well as the potential impact of stratospheric ozone depletion on global ecosystems. As both photosynthesis and UV fluxes are subject to diurnal variations, we examined the diurnal variability of the effect of UV radiation on photosynthesis in three diverse algal mats. In one of the mats (Cyanidium caldarium) a small mean decrease in primary productivity over the whole day occurred when both UVA and UVB were screened out. In two of the mats (Lyngbya aestuarii and Zygogonium sp.) we found a mean increase in the total primary productivity over the day when UVB alone was screened and a further increase when UVA and UVB were both screened out. Variations in the effects of UV radiation were found at different times of the day. This diurnal variability may be because even under the same solar radiation flux, there are different factors that may control photosynthetic rate, including nutritional status and other physiological processes in the cell. The results show the importance of assessing the complete diurnal productivity. For some of the time points the increase in the mean was still within the standard deviations in primary productivity, illustrating the difficulty in dissecting UV effects from other natural variations.

Ultraviolet Rays↗

NASA NTRS2003

Real-Time Imaging of Ground Cover: Relationships with Radiation Capture, Canopy Photosynthesis, and Daily Growth Rate

Cumulative absorbed radiation is highly correlated with crop biomass and yield. In this chapter we describe the use of a digital camera and commercial imaging software for estimating daily radiation capture, canopy photosynthesis, and relative growth rate. Digital images were used to determine percentage of ground cover of lettuce (Lactuca sativa L.) communities grown at five temperatures. Plants were grown in a steady-state, 10-chamber CO2 gas exchange system, which was used to measure canopy photosynthesis and daily carbon gain. Daily measurements of percentage of ground cover were highly correlated with daily measurements of both absorbed radiation (r(sup 2) = 0.99) and daily carbon gain (r(sup 2) = 0.99). Differences among temperature treatments indicated that these relationships were influenced by leaf angle, leaf area index, and chlorophyll content. An analysis of the daily images also provided good estimates of relative growth rates, which were verified by gas exchange measurements of daily carbon gain. In a separate study we found that images taken at hourly intervals were effective for monitoring real-time growth. Our data suggests that hourly images can be used for early detection of plant stress. Applications, limitations, and potential errors are discussed. We have long known that crop yield is determined by the efficiency of four component processes: (i) radiation capture, (ii) quantum yield, (iii) carbon use efficiency, and (iv) carbon partitioning efficiency (Charles-Edwards, 1982; Penning de Vries & van Laar, 1982; Thornley, 1976). More than one-half century ago, Watson (1947, 1952) showed that variation in radiation capture accounted for almost all of the variation in yield between sites in temperate regions, because the three other components are relatively constant when the crop is not severely stressed. More recently, Monteith (1977) reviewed the literature on the close correlation between radiation capture and yield. Bugbee and Monje (1992) demonstrated the close relationship between absorbed radiation and yield in an optimal environment.

Klassen, S. P.↗

NASA NTRS2005

Acclimation of Plant Populations to Shade: Photosynthesis, Respiration, and Carbon Use Efficiency

Cloudy days cause an abrupt reduction in daily photosynthetic photon flux (PPF), but we have a poor understanding of how plants acclimate to this change. We used a unique lo-chamber, steady-state, gas-exchange system to continuously measure daily photosynthesis and night respiration of populations of a starch accumulator [tomato (Lycopersicone scukntum Mill. cv. Micro-Tina)] and a sucrose accumulator [lettuce (Latuca sativa L ev. Grand Rapids)] over 42 days. AI1 measurements were done at elevated CO2, (1200micr-/mol) avoid any CO2 limitations and included both shoots and roots. We integrated photosynthesis and respiration measurements separately to determine daily net carbon gain and carbon use efficiency (CUE) as the ratio of daily net C gain to total day-time C fixed over the 42-day period. After 16 to 20 days of growth in constant PPF, plants in some chambers were subjected to an abrupt PPF reduction to simulate shade or a series of cloudy days. The immediate effect and the long term acclimation rate w'ere assessed from canopy quantum yield and carbon use efficiency. The effect of shade on carbon use efficiency and acclimation was much slower than predicted by widely used growth models. It took 12 days for tomato populations to recover their original CUE and lettuce CUE never completely acclimated. Tomatoes, the starch accumulator, acclimated to low light more rapidly than lettuce, the sucrose accumulator. Plant growth models should be modified to include the photosynthesis/respiration imbalance and resulting inefficiency of carbon gain associated with changing PIT conditions on cloudy days.

Frantz, Jonathan M.↗

NASA NTRS2009

High-Efficiency Artificial Photosynthesis Using a Novel Alkaline Membrane Cell

A new cell designed to mimic the photosynthetic processes of plants to convert carbon dioxide into carbonaceous products and oxygen at high efficiency, has an improved configuration using a polymer membrane electrolyte and an alkaline medium. This increases efficiency of the artificial photosynthetic process, achieves high conversion rates, permits the use of inexpensive catalysts, and widens the range of products generated by this type of process. The alkaline membrane electrolyte allows for the continuous generation of sodium formate without the need for any additional separation system. The electrolyte type, pH, electrocatalyst type, and cell voltage were found to have a strong effect on the efficiency of conversion of carbon dioxide to formate. Indium electrodes were found to have higher conversion efficiency compared to lead. Bicarbonate electrolyte offers higher conversion efficiency and higher rates than water solutions saturated with carbon dioxide. pH values between 8 and 9 lead to the maximum values of efficiency. The operating cell voltage of 2.5 V, or higher, ensures conversion of the carbon dioxide to formate, although the hydrogen evolution reaction begins to compete strongly with the formate production reaction at higher cell voltages. Formate is produced at indium and lead electrodes at a conversion efficiency of 48 mg of CO2/kilojoule of energy input. This efficiency is about eight times that of natural photosynthesis in green plants. The electrochemical method of artificial photosynthesis is a promising approach for the conversion, separation and sequestration of carbon dioxide for confined environments as in space habitats, and also for carbon dioxide management in the terrestrial context. The heart of the reactor is a membrane cell fabricated from an alkaline polymer electrolyte membrane and catalyst- coated electrodes. This cell is assembled and held in compression in gold-plated hardware. The cathode side of the cell is supplied with carbon dioxide-saturated water or bicarbonate solution. The anode side of the cell is supplied with sodium hydroxide solution. The solutions are circulated past the electrodes in the electrochemical cell using pumps. A regulated power supply provides the electrical energy required for the reactions. Photovoltaic cells can be used to better mimic the photosynthetic reaction. The current flowing through the electrochemical cell, and the cell voltage, are monitored during experimentation. The products of the electrochemical reduction of carbon dioxide are allowed to accumulate in the cathode reservoir. Samples of the cathode solution are withdrawn for product analysis. Oxygen is generated on the anode side and is allowed to vent out of the reservoir.

Narayan, Sri↗

NASA NTRS2012

Effects of CO2 Concentration on Leaf Photosynthesis and Stomatal Conductance of Potatoes Grown Under Different Irradiance Levels and Photoperiods

Potato (Solanum tuberosum L.) cvs. Russet Burbank, Denali, and Norland, were grown in environmental rooms controlled at approx 350 micro mol/mol (ambient during years 1987/1988) and 1000 micro mol/mol (enriched) CO2 concentrations. Plants and electric lamps were arranged to provide two irradiance zones, 400 and 800 micro mol/mol/square m/S PPF and studies were repeated using two photoperiods (12-h light / 12-h dark and continuous light). Leaf photosynthetic rates and leaf stomatal conductance were measured using fully expanded, upper canopy leaves at weekly intervals throughout growth (21 through 84 days after transplanting). Increasing the CO2 from approx 350 to 1000 micro mol/mol under the 12-h photoperiod increased leaf photosynthetic rates by 39% at 400 micro mol/mol/square m/S PPF and 27% at 800 micro mol/mol/square m/S PPF. Increasing the CO2 from approx 350 to 1000 micro mol/mol under continuous light decreased leaf photosynthetic rates by 7% at 400 micro mol/mol/square m/S PPF and 13% at 800 micro mol/mol/square m/S PPF. Increasing the CO2 from approx 350 to 1000 micro mol/mol under the 12-h photoperiod plants decreased stomatal conductance by an average of 26% at 400 micro mol/mol/square m/S PPF and 42% at 800 micro mol/mol/square m/S PPF. Under continuous light, CO2 enrichment resulted in a small increase (2%) of stomatal conductance at 400 micro mol/mol/square m/S PPF, and a small decrease (3%) at 800 micro mol/mol/square m/S PPF. Results indicate that CO2 enrichment under the 12-h photoperiod showed the expected increase in photosynthesis and decrease in stomatal conductance for a C3 species like potato, but the decreases in leaf photosynthetic rates and minimal effect on conductance from CO2 enrichment under continuous light were not expected. The plant leaves under continuous light showed more chlorosis and some rusty flecking versus plants under the 12-h photoperiod, suggesting the continuous light was more stressful on the plants. The increased rates of leaf photosynthesis with increased CO2 concentration paralleled trends in biomass production (published previously) but were not proportional to the biomass yields.

Wheeler, R. M.↗

NASA NTRS2011

Efficiency of Photosynthesis in a Chl d-Utilizing Cyanobacterium is Comparable to or Higher than that in Chl a-Utilizing Oxygenic Species

The cyanobacterium Acaryochloris marina uses chlorophyll d to carry out oxygenic photosynthesis in environments depleted in visible and enhanced in lower-energy, far-red light. However, the extent to which low photon energies limit the efficiency of oxygenic photochemistry in A. marina is not known. Here, we report the first direct measurements of the energy-storage efficiency of the photosynthetic light reactions in A. marina whole cells,and find it is comparable to or higher than that in typical, chlorophyll a-utilizing oxygenic species. This finding indicates that oxygenic photosynthesis is not fundamentally limited at the photon energies employed by A. marina, and therefore is potentially viable in even longer-wavelength light environments.

Mielke, S. P.↗