Photosynthetic Energy Conversion Efficiency Research Articles

Photosynthetic energy conversion efficiency: setting a baseline for gauging future improvements in important food and biofuel crops.

The conversion efficiency (ε(c)) of absorbed radiation into biomass (MJ of dry matter per MJ of absorbed photosynthetically active radiation) is a component of yield potential that has been estimated at less than half the theoretical maximum. Various strategies have been proposed to improve ε(c), but a statistical analysis to establish baseline ε(c) levels across different crop functional types is lacking. Data from 164 published ε(c) studies conducted in relatively unstressed growth conditions were used to determine the means, greatest contributors to variation, and genetic trends in ε(c )across important food and biofuel crop species. ε(c) was greatest in biofuel crops (0.049-0.066), followed by C4 food crops (0.046-0.049), C3 nonlegumes (0.036-0.041), and finally C3 legumes (0.028-0.035). Despite confining our analysis to relatively unstressed growth conditions, total incident solar radiation and average growing season temperature most often accounted for the largest portion of ε(c) variability. Genetic improvements in ε(c), when present, were less than 0.7% per year, revealing the unrealized potential of improving ε(c) as a promising contributing strategy to meet projected future agricultural demand.

Structure and energy transfer in photosystems of oxygenic photosynthesis.

Oxygenic photosynthesis is the principal converter of sunlight into chemical energy on Earth. Cyanobacteria and plants provide the oxygen, food, fuel, fibers, and platform chemicals for life on Earth. The conversion of solar energy into chemical energy is catalyzed by two multisubunit membrane protein complexes, photosystem I (PSI) and photosystem II (PSII). Light is absorbed by the pigment cofactors, and excitation energy is transferred among the antennae pigments and converted into chemical energy at very high efficiency. Oxygenic photosynthesis has existed for more than three billion years, during which its molecular machinery was perfected to minimize wasteful reactions. Light excitation transfer and singlet trapping won over fluorescence, radiation-less decay, and triplet formation. Photosynthetic reaction centers operate in organisms ranging from bacteria to higher plants. They are all evolutionarily linked. The crystal structure determination of photosynthetic protein complexes sheds light on the various partial reactions and explains how they are protected against wasteful pathways and why their function is robust. This review discusses the efficiency of photosynthetic solar energy conversion.

Cold acclimation and BnCBF17-over-expression enhance photosynthetic performance and energy conversion efficiency during long-term growth of Brassica napus under elevated CO2 conditions

The effects of cold acclimation and long-term elevated CO(2) on photosynthetic performance of wild-type (WT) and BnCBF17-over-expressing line of Brassica napus cv. Westar (BnCBF17-OE) grown at either 20/16°C (non-acclimated) or 5/5°C (cold acclimated) and at either ambient (380μmolCmol(-1)) or elevated (700μmolCmol(-1)) CO(2) were studied. Compared with non-acclimated WT, the BnCBF17-OE grown at 20°C mimicked the effects of cold acclimation on WT B. napus with respect to compact dwarf phenotype and increased rates of light-saturated CO(2) assimilation and photosynthetic electron transport. This was associated with enhanced energy conversion efficiency into biomass as assessed by decreased excitation pressure coupled to decreased dependence on non-photochemical energy dissipation for a given irradiance. Growth at elevated CO(2) decreased the light and CO(2)-saturated rates of photosynthesis by 30% for non-acclimated WT relative to growth at ambient CO(2). This was associated with inhibition in electron transport rates (20%), decrease in amount of rbcL (35%) and cytosolic FBPase (70%) and increased excitation pressure and non-photochemical quenching in elevated versus ambient CO(2)-grown non-acclimated WT. In contrast, light and CO(2)-saturated rates of photosynthesis, electron transport, excitation pressure, non-photochemical quenching and levels of rbcL, cytosolic FBPase and Lhcb1 were insensitive to growth under elevated CO(2) in BnCBF17-OE and cold-acclimated WT. Thus, BnCBF17-over-expression and cold acclimation maintain enhanced energy conversion efficiency and reduced sensitivity to feedback-limited photosynthesis during long-term growth of B. napus under elevated CO(2). Our results indicated that CBFs transcription factors regulate not only freezing tolerance but also has major whole plant effects.

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

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 3,000m). 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.

Improving photosynthesis for algal biofuels: toward a green revolution

Biofuels derived from marine algae are a potential source of sustainable energy that can contribute to future global demands. The realisation of this potential will require manipulation of the fundamental biology of algal physiology to increase the efficiency with which solar energy is ultimately converted into usable biomass. This 'photosynthetic solar energy conversion efficiency' sets an upper limit on the potential of algal-derived biofuels. In this review, we outline photosynthetic molecular targets that could be manipulated to increase the efficiency and yield of algal biofuel production. We also highlight modern 'omic' and high-throughput technologies that might enable identification, selection and improvement of algal cell lines on timescales relevant for achieving significant contributions to future energy solutions.

Expression of yeast Hem1 controlled by Arabidopsis HemA1 promoter enhances leaf photosynthesis in transgenic tobacco

A gene encoding aminolevulinate synthase (ALA-S) in yeast (Saccharomyces cerevisiae YHem1) was introduced into the genome of tobacco (Nicoliana tabacum) under the control of Arabidopsis thaliana HemA1 gene promoter (AtHemA1 P). All transgenic lines transcribed the YHem1 gene, especially under light condition. The capacity to synthesize ALA and therefore chlorophyll was increased in transformed plants. Determination of gas exchange suggested that transgenic plants had significantly higher level of net photosynthetic rate (P ( n )), stomatal conductance (G ( s )) and transpiration rate (T ( r )), compared to the wild type (WT). Analysis with a modulated chlorophyll fluorometer demonstrated that the genetic transformation also caused a significant increase in photochemical efficiency of PSII ([Formula: see text]), actual photochemical efficiency (Ф ( PSII )), photochemical quenching (qP), electron transfer rate (ETR) and the energy proportion in photochemistry (Pc), but decrease in proportion in heat dissipation (Hd). Chlorophyll-a fast fluorescence measurement and JIP-test indicated that photosynthetic performance index on cross section basis (PI ( CS )) and electron transport flux (ET ( o ) /CS) of transgenic tobacco were increased remarkably. And the probability that a trapped exciton can move a electron into the electron transport chain beyond Q ( A ) (-) (Ψ ( o )) and the density of active reaction centers (RC/CS) were also increased obviously in transgenic tobacco. These results imply that transgenic tobacco plants expressing YHem1 gene had higher photosynthetic capacity and energy conversion efficiency than the WT plants.

Systems-level modeling—A new approach for engineering efficient photosynthetic machinery

Photosynthesis produces the basic building block for crop biomass and yields; however, improving photosynthesis has not been effectively used as a breeding goal. More and more evidences suggested that improving photosynthesis can substantially increase crop yields. The complexity of photosynthesis however makes experimentally identifying new ways to engineer higher photosynthesis inherently time-consuming and costly. Combining systems modeling with evolutionary algorithm makes it possible to identify optimal engineering options for future global climate change scenarios and simultaneously consider environmental constraints, such as with constant or even decreasing nitrogen fertilizer application in the field. This method enables in silico examination of a large number of engineering options which natural selection has not explored, for higher photosynthetic energy conversion efficiency. The new approach comes particularly timely for now when our society is facing serious challenges in food security and global climate change. The traditional reductionist's approach will continue generating critical knowledge required to support this systems biology method to engineering higher photosynthesis.

Linear dichroism and circular dichroism in photosynthesis research.

The efficiency of photosynthetic light energy conversion depends largely on the molecular architecture of the photosynthetic membranes. Linear- and circular-dichroism (LD and CD) studies have contributed significantly to our knowledge of the molecular organization of pigment systems at different levels of complexity, in pigment–protein complexes, supercomplexes, and their macroassemblies, as well as in entire membranes and membrane systems. Many examples show that LD and CD data are in good agreement with structural data; hence, these spectroscopic tools serve as the basis for linking the structure of photosynthetic pigment–protein complexes to steady-state and time-resolved spectroscopy. They are also indispensable for identifying conformations and interactions in native environments, and for monitoring reorganizations during photosynthetic functions, and are important in characterizing reconstituted and artificially constructed systems. This educational review explains, in simple terms, the basic physical principles, and theory and practice of LD and CD spectroscopies and of some related quantities in the areas of differential polarization spectroscopy and microscopy.Electronic supplementary materialThe online version of this article (doi:10.1007/s11120-009-9424-4) contains supplementary material, which is available to authorized users.

What is the maximum efficiency with which photosynthesis can convert solar energy into biomass?

Photosynthesis is the source of our food and fiber. Increasing world population, economic development, and diminishing land resources forecast that a doubling of productivity is critical in meeting agricultural demand before the end of this century. A starting point for evaluating the global potential to meet this goal is establishing the maximum efficiency of photosynthetic solar energy conversion. The potential efficiency of each step of the photosynthetic process from light capture to carbohydrate synthesis is examined. This reveals the maximum conversion efficiency of solar energy to biomass is 4.6% for C3 photosynthesis at 30 degrees C and today's 380 ppm atmospheric [CO2], but 6% for C4 photosynthesis. This advantage over C3 will disappear as atmospheric [CO2] nears 700 ppm.

An in situ iron-enrichment experiment in the western subarctic Pacific (SEEDS): Introduction and summary

To test the iron hypothesis in the subarctic Pacific Ocean, an in situ iron-enrichment experiment (SEEDS) was performed in the western subarctic gyre in July–August 2001. About 350 kg of iron (as acidic iron sulfate) and 0.48 mol of the inert chemical tracer sulfur hexafluoride were introduced into a 10-m deep surface mixed layer over an 80 km 2 area. This single iron infusion raised dissolved iron levels to ∼2.9 nM initially. Dissolved iron concentrations rapidly decreased after the infusion, but levels remained close to 0.15 nM even at the end of the 14-day experimental period. During SEEDS there were iron-mediated increases in chlorophyll a concentrations (up to 20 μg l −1), primary production rates, biomass and photosynthetic energy conversion efficiency relative to waters outside the iron-enriched patch. The rapid and very high accumulation of phytoplankton biomass in response to the iron addition appeared to be partly attributable to shallow mixed-layer depth and moderate water temperature in the western subarctic Pacific. However, the main reason was a floristic shift to fast-growing centric diatom Chaetoceros debilis, unlike the previous iron-enrichment experiments in the equatorial Pacific and the Southern Ocean, in both of which iron stimulated the growth of pennate diatoms. The iron-mediated blooming of diatoms resulted in a marked consumption of macronutrients and drawdown of pCO 2. Biological and physiological measurements indicate that phytoplankton growth in the patch became both light- and iron-limited, making phytoplankton biomass relatively constant after day 9. The increase in microzooplankton grazing rate after day 9 also influenced the net growth rate of phytoplankton. There was no significant increase in the export flux of carbon to depth during the 14-day occupation of the experimental site. The export flux between day 4 and day 13 was estimated to be only 13% of the integrated primary production in the iron-enriched patch. The major part of the carbon fixed by the diatom bloom remained in the surface mixed layer as biogenic particulate matter. Our findings support the hypothesis that iron limits phytoplankton growth and biomass in a ‘bottom up’ manner in this area, but the fate of algal carbon remains unknown.

Effect of difference in nitrogen media on salt-stress response and contents of nitrogen compounds in rice seedlings

Effect of the difference in nitrogen media on the salt stress response and contents of nitrogen compounds (amino acids, polyamines, and protein) was investigated in rice seedlings (Oryza sativa L. cv. Nipponbare) at the 2nd leaf stage. Plant growth and chlorophyll fluorescence (ΦII), reflecting the efficiency of photosynthetic energy conversion, markedly decreased during the 3-d period of the 100 mM NaCl treatment. Response to salt stress, however, varied with the difference in the nitrogen media. Rice seedlings grown in a low ammonium medium, containing a sub-optimal concentration of ammonium for seedling growth, showed the lowest sensitivity to the salt stress treatment. In these seedlings, free amino acids, especially amide glutamine and arginine, strikingly accumulated in the 2nd leaf blades with the NaCl treatment. Additionally, the leaf polyamine contents in the seedlings grown in a low ammonium medium increased under salt stress, while the leaf polyamine contents in the seedlings grown in high ammonium and nitrate media considerably decreased. Under salt stress conditions, a positive correlation was observed between the total polyamine content and ΦII in the 2nd leaf blades. These results suggest that the salt stress tolerance of rice seedlings is associated with effective assimilation of excess ammonia, induced by salt stress, by nitrogen compounds such as arginine, glutamine, and polyamines.

EFFECTS OF UV RADIATION ON A CHLOROPHYTE ALGA (SCENEDESMUS SP.) ISOLATED FROM THE FUMAROLE FIELDS OF MT. EREBUS, ANTARCTICA1

Acclimation to UV radiation (UVR) was examined in a unicellular chlorophyte isolated from fumarole fields adjacent to Mt. Erebus, Antarctica. Long‐term exposure to UVR (14 days) initially reduced the efficiency of photosynthetic energy conversion measured as the dark adapted quantum yield of PSII fluorescence (Fv/Fm) when compared with cultures not exposed to UVR. However, the UVR exposed cultures recovered to pre‐UVR exposure levels of photosynthetic efficiency by the end of the experimental period. Additionally, neither growth rates nor cell size were significantly affected by exposure to UVR. In contrast, the maximum rate of carbon fixation on a chl a basis was significantly reduced at the end of the experiment and was accompanied by a significant decrease in chl a concentrations. These results suggest a transient effect of UVR on primary photochemistry consistent with damage to PSII, possibly at the D1 protein, with more chronic effects on carbon fixation that did affect maximum photosynthetic capacity. Ultrastructural and molecular (18s rDNA) data show that this isolate from the fumoral fields of Mt. Erebus, Antarctica is a member of the genus Scenedesmus.

NEW EMBO MEMBER'S REVIEW: State transitions reveal the dynamics and flexibility of the photosynthetic apparatus

The chloroplast-based photosynthetic apparatus of plants and algae associates various redox cofactors and pigments with approximately 70 polypeptides to form five major transmembrane protein complexes. Among these are two photosystems that have distinct light absorption properties but work in series to produce reducing equivalents aimed at the fixation of atmospheric carbon. A short term chromatic adaptation known as 'State transitions' was discovered thirty years ago that allows photosynthetic organisms to adapt to changes in light quality and intensity which would otherwise compromise the efficiency of photosynthetic energy conversion. A two-decade research effort has finally unraveled the major aspects of the molecular mechanism responsible for State transitions, and their physiological significance has been revisited. This review describes how a-still elusive-regulatory kinase senses the physiological state of the photosynthetic cell and triggers an extensive supramolecular reorganization of the photosynthetic membranes. The resulting picture of the photosynthetic apparatus is that of a highly flexible energy convertor that adapts to the ever-changing intracellular demand for ATP and/or reducing power.

Effects of zinc and influence of Acremonium lolii on growth parameters, chlorophyll a fluorescence and antioxidant enzyme activities of ryegrass (Lolium perenne L. cv Apollo)

The effects of zinc on growth, mineral content, chlorophyll a fluorescence, and detoxifying enzyme activity (ascorbate peroxidase (APX), EC 1.11.1.11; superoxide dismutase (SOD), EC 1.15.1.1) of ryegrass infected or not by Acremonium lolii, and treated with nutrient solution containing 0-50 mM ZnSO(4) were studied. The introduction of zinc induces stress with a decrease in growth at 1, 5 and 10 mM ZnSO(4) and a cessation of growth at 50 mM ZnSO(4), in ryegrass plants infected by A. lolii or not. This decrease in growth may be due to an accumulation of zinc in leaves. Nevertheless, symbiotic plants showed higher values in tiller number, an advantage conferred by the fungus. After 24 d of Zn exposure, leaf fresh weights and leaf water content were lower in plants growing with Zn in the culture medium and no advantage was conferred by the fungus to its host. An increase in Zn supply resulted in a decrease of the Ca, K, Mg, and Cu content of the leaves, a reduction in the quantum yield of electron flow throughout photosystem II (DeltaF/F(1)(m))and a lowering of the efficiency of photosynthetic energy conversion (F(v)/F(m)), compared to control plants. To counter this zinc stress, detoxifying enzymes APX and SOD increased (100%) when Zn reached the value of 50 mM in the nutrient solution. At 10 mM ZnSO(4), the presence of the fungus in the plant led to an increase in the threshold toxicity of plants to zinc by a diminution of APX activity.

IronEx-I, an in situ iron-enrichment experiment: Experimental design, implementation and results

An in situ iron-enrichment experiment near the Galapagos Islands was performed in October 1993. Here we report the theoretical and practical considerations of creating such a patch of iron-enriched surface water, as well as the strategies employed for the detection of the patch and the biological and chemical signals which developed, in an area dominated by advective processes. Physical and chemical models were used to predict the speciation, solubility, and the final concentration of iron in surface waters injected with acidic iron sulfate. A trial injection off the California coast in which 800 L of a 0.5 M FeSO 4 were introduced into the ship’s wake over a 1.5 km 2 area, was used to test these predictions. Iron concentrations were determined continually onboard during the initial experiment as the ship steamed in transects through the enriched patch. The results indicate excellent spatial agreement with model predictions and final concentrations that were consistent with the chemical model. However, the use of a Cartesian coordinate system during injection resulted in an extremely compressed, heterogeneous patch. Results from this preliminary experiment were then applied towards the development and implementation of the first open ocean iron enrichment experiment (IronEx I) near the Galapagos Islands in October 1993. The development and results of these methodologies are presented. In the IronEx I equatorial experiment, a Lagrangian coordinate system was established using a drogued buoy (equipped with GPS and packet radio) and the iron-enriched area (64 km 2 containing 443 kg of Fe) was tagged with the inert chemical tracer sulfurhexafluoride (SF 6). This strategy resulted in a fairly rectangular, homogeneous enriched patch initially detectable by both Fe and SF 6 determination. Shipboard analysis and airborne observations confirmed good spatial agreement between the Lagrangian drifter and the biological and chemical signatures in the patch. Biological and chemical sampling of the enriched area showed an increase in chlorophyll, primary production, biomass and photosynthetic energy conversion efficiency relative to waters outside the patch, supporting the hypothesis that iron limits phytoplankton growth and biomass in a ‘bottom up’ manner in this area. The ability to create a coherent patch and track it over time led to this first open-ocean test of the iron hypothesis.

PERSPECTIVES IN COMPARATIVE ECOPHYSIOLOGY OF SOME BRAZILIAN VEGETATION TYPES: LEAF CO2 ANDH2O GAS EXCHANGE, CHLOROPHYLL a FLUORESCENCE AND CARBON ISOTOPE DISCRIMINATION

Photosynthetic activity is responsible for the energy plants use for growth, defense and reproduction. Therefore, photosynthesis is fundamental to plant metabolism However, the effects of environmental factors on net CO 2 assimilation rates are as yet poorly known in Brazilian vegetation types. A decrease in net CO 2 uptake may be related to a decrease in CO, diffusion to the carboxylation sites due to partial or total stomata closure and to direct inhibition 01' primary and secondary photosynthetic processes in the chloroplasts. Photosynthetic carbon reduction is one of the rnost important pathways for the utilization of light energy. Hence, lower rates of CO, uptake may cause an excess of excitation energy through the photosystems and consequently le ad to a decline in the efficiency of photosynthetic energy conversion. The study of comparative ecophysiology of native species is essential for understanding the mechanisms that lead to the high diversity of the Brazilian flora. CO, and H 2 O gas exchange, chlorophyll a fluorescence and carbon isotope discrimination are powerful and predictive tools for addressing the influence of multi pie environmental factors on plant performance in situ. Here 1 discuss the results of comparative plant physiological ecology studies in several Brazilian vegetation types.

PERSPECTIVES IN COMPARATIVE ECOPHYSIOLOGY OF SOME BRAZILIAN VEGETATION TYPES: LEAF CO2 AND H2O GAS EXCHANGE, CHLOROPHYLL A FLUORESCENCE AND CARBON ISOTOPE DISCRIMINATION

Photosynthetic activity is responsible for the energy plants use for growth, defense and reproduction. Therefore, photosynthesis is fundamental to plant metabolism However, the effects of environmental factors on net CO 2 assimilation rates are as yet poorly known in Brazilian vegetation types. A decrease in net CO 2 uptake may be related to a decrease in CO, diffusion to the carboxylation sites due to partial or total stomata closure and to direct inhibition 01' primary and secondary photosynthetic processes in the chloroplasts. Photosynthetic carbon reduction is one of the rnost important pathways for the utilization of light energy. Hence, lower rates of CO, uptake may cause an excess of excitation energy through the photosystems and consequently le ad to a decline in the efficiency of photosynthetic energy conversion. The study of comparative ecophysiology of native species is essential for understanding the mechanisms that lead to the high diversity of the Brazilian flora. CO, and H 2 O gas exchange, chlorophyll a fluorescence and carbon isotope discrimination are powerful and predictive tools for addressing the influence of multi pie environmental factors on plant performance in situ. Here 1 discuss the results of comparative plant physiological ecology studies in several Brazilian vegetation types.

Effects of photoinhibitory treatment on CO2 assimilation, the quantum yield of CO2 assimilation, D1 protein, ascorbate, glutathione and xanthophyll contents and the electron transport rate in vine leaves

ABSTRACTThe responses of photosynthesis to high light and low temperature were studied in vines cultivated in the greenhouse in low light. Exposure to high light (1000 /umol m−2 s−1) or low temperature (5 °C) alone had no measurable effect on the photosynthetic processes, but the combination of high light and low temperature caused rapid loss of photosynthetic capacity and a decrease in the efficiency of photosynthetic energy conversion. After a 15 h exposure to 5°C at high light, the Fv/sb/Fmratio had decreased by 80% and the apparent quantum yield by 75%. Nevertheless, when the leaves were returned to low light at 22°C, these parameters recovered rapidly. The foliar pools of ascorbate and glutathione decreased in the first hours of photoinhibitory treatment while the zeaxanthin content increased from negligible levels to about 50% of the total foliar xanthophyll pool. There was a clear correlation between the zeaxanthin content of the leaves and their Fv/Fm ratio during both photoinhibition and recovery. However, there was also a good correlation between the decrease in theFv Fm ratio and the measured decrease in the total foliar levels of the antioxidants ascorbate and glutathione. The amount of D, protein diminished over the same period as the zeaxanthin levels were increasing. This approach, involving simultaneous measurements of several parameters considered to influence photosystemy II activity, clearly demonstrates that measured decreases in Fv/Fm may not simply be related to zeaxanthin levels or to amounts of D1 protein alone but result from multifactoral influences.

Variations in Chlorophyll Fluorescence Yields in Phytoplankton in the World Oceans

The ocean is optically thin and lends itself to large-scale measurements of in vivo chlorophyll fluorescence. In the open ocean, however, phytoplankton chlorophyll concentrations average only 0.2 μg L-1, and hence high sensitivity is required for precise measurements of the fluorescence yields. Over the past decade, we have developed two approaches to achieve the required sensitivity; these are the pump- and probe-technique and a fast repetition rate (FRR) method. Both methods have been adapted for in situ studies and are used to rapidly measure the maximum change in the quantum yield (Δ�max) of photosystem II (PSII), as well as the effective absorption cross-section of PSII (σPSII). Sections of variable fluorescence across the Pacific and Atlantic Oceans reveal the influence of geophysical processes in controlling the quantum yields of phytoplankton photosynthesis. Areas of upwelling, such as off the coast of north-westem Africa, have Fv/Fm values of 0.65, which are close to the maximum achievable values in nutrient-replete cultures. Throughout most of the nutrient-deficient central ocean basins, this quantum efficiency is reduced by more than 50%. In high-nutrient, low- chlorophyll regions of the eastern Equatorial Pacific, the deliberate, large-scale addition of nanomolar iron directly to the ocean leads to a rapid increase in quantum efficiency of the natural phytoplankton community, thereby revealing that in these regions phytoplankton photosynthetic energy conversion efficiency is iron limited. Diel patterns of variation in the upper ocean display midday, intensity- dependent reductions in both upsII and A�max. We interpret the former as indicative of non- photochemical quenching in the antenna, while the latter is a consequence of both rapidly reversible and slowly reversible damage to reaction centres. From knowledge of the incident spectral irradiance, Δ�max, σPSII, and photochemical quenching, the absolute photosynthetic electron transport rate can be derived in real-time. Using unattended, moored continuous measurements of in vivo fluorescence parameters, the derived in situ electron transport rates can be related to satellite observations of the global ocean with basin-scale, seasonal estimates of phytoplankton carbon fixation. Thus, unlike any other photosynthetic parameter, chlorophyll fluorescence can be used to bridge the scales of biophysical responses to ecosystem dynamics.

Iron-Induced Changes in Light Harvesting and Photochemical Energy Conversion Processes in Eukaryotic Marine Algae

The role of iron in regulating light harvesting and photochemical energy conversion processes was examined in the marine unicellular chlorophyte Dunaliella tertiolecta and the marine diatom Phaeodactylum tricornutum. In both species, iron limitation led to a reduction in cellular chlorophyll concentrations, but an increase in the in vivo, chlorophyll-specific, optical absorption cross-sections. Moreover, the absorption cross-section of photosystem II, a measure of the photon target area of the traps, was higher in iron-limited cells and decreased rapidly following iron addition. Iron-limited cells exhibited reduced variable/maximum fluorescence ratios and a reduced fluorescence per unit absorption at all wave-lengths between 400 and 575 nm. Following iron addition, variable/maximum fluorescence ratios increased rapidly, reaching 90% of the maximum within 18 to 25 h. Thus, although more light was absorbed per unit of chlorophyll, iron limitation reduced the transfer efficiency of excitation energy in photosystem II. The half-time for the oxidation of primary electron acceptor of photosystem II, calculated from the kinetics of decay of variable maximum fluorescence, increased 2-fold under iron limitation. Quantitative analysis of western blots revealed that cytochrome f and subunit IV (the plastoquinone-docking protein) of the cytochrome b(6)/f complex were also significantly reduced by lack of iron; recovery from iron limitation was completely inhibited by either cycloheximide or chloramphenicol. The recovery of maximum photosynthetic energy conversion efficiency occurs in three stages: (a) a rapid (3-5 h) increase in electron transfer rates on the acceptor side of photosystem II correlated with de novo synthesis of the cytochrome b(6)/f complex; (b) an increase (10-15 h) in the quantum efficiency correlated with an increase in D1 accumulation; and (c) a slow (>18 h) increase in chlorophyll levels accompanied by an increase in the efficiency of energy transfer from the light-harvesting chlorophyll proteins to the reaction centers.