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Molecular evidence for the early evolution of photosynthesis.

The origin and evolution of photosynthesis have long remained enigmatic due to a lack of sequence information of photosynthesis genes across the entire photosynthetic domain. To probe early evolutionary history of photosynthesis, we obtained new sequence information of a number of photosynthesis genes from the green sulfur bacterium Chlorobium tepidum and the green nonsulfur bacterium Chloroflexus aurantiacus. A total of 31 open reading frames that encode enzymes involved in bacteriochlorophyll/porphyrin biosynthesis, carotenoid biosynthesis, and photosynthetic electron transfer were identified in about 100 kilobase pairs of genomic sequence. Phylogenetic analyses of multiple magnesium-tetrapyrrole biosynthesis genes using a combination of distance, maximum parsimony, and maximum likelihood methods indicate that heliobacteria are closest to the last common ancestor of all oxygenic photosynthetic lineages and that green sulfur bacteria and green nonsulfur bacteria are each other's closest relatives. Parsimony and distance analyses further identify purple bacteria as the earliest emerging photosynthetic lineage. These results challenge previous conclusions based on 16S ribosomal RNA and Hsp60/Hsp70 analyses that green nonsulfur bacteria or heliobacteria are the earliest phototrophs. The overall consensus of our phylogenetic analysis, that bacteriochlorophyll biosynthesis evolved before chlorophyll biosynthesis, also argues against the long-held Granick hypothesis.

Bacteria↗

Effect of mercuric ion on the growth, photosynthesis, and nitrogenase activity of Anabaena inaequalis.

Anabaena inaequalis was sensitive to mercuric ion (Hg2+) in the ppb (nanogram per milliliter) range. Growth was inhibited significantly at concentrations of metal ion as low as 2 ppb, and 100 ppb was required to inhibit photosynthesis and acetylene reduction. Low levels of Hg2+ stimulated acetylene reduction and photosynthesis. The lysis of vegetative cells was the primary action of mercuric ions, resulting in the inhibition of growth, photosynthesis, and nitrogenase activity. There was a linear relationship between numbers of cells and the amount of Hg2+ required to induce culture lysis. Calculated on the basis of equivalent cell numbers, Hg2+ was toxic to A. inaequalis at 0.006, 0.009, and 0.100 micrograms of Hg2+ per 10(5) cells for photosynthesis, growth, and acetylene reduction, respectively.

Acetylene↗

Complex evolution of photosynthesis.

The origin of photosynthesis is a fundamental biological question that has eluded researchers for decades. The complexity of the origin and evolution of photosynthesis is a result of multiple photosynthetic components having independent evolutionary pathways. Indeed, evolutionary scenarios have been established for only a few photosynthetic components. Phylogenetic analysis of Mg-tetrapyrrole biosynthesis genes indicates that most anoxygenic photosynthetic organisms are ancestral to oxygen-evolving cyanobacteria and that the purple bacterial lineage may contain the most ancestral form of this pigment biosynthesis pathway. The evolutionary path of type I and type II reaction center apoproteins is still unresolved owing to the fact that a unified evolutionary tree cannot be generated for these divergent reaction center subunits. However, evidence for a cytochrome b origin for the type II reaction center apoproteins is emerging. Based on the combined information for both photopigments and reaction centers, a unified theory for the evolution of reaction center holoproteins is provided. Further insight into the evolution of photosynthesis will have to rely on additional broader sampling of photosynthesis genes from divergent photosynthetic bacteria.

Amino Acid Sequence↗

Expression pattern of Stlhcb gene family in potato and effects of overexpression of Stcp24 gene on potato photosynthesis.

Potato is one of the four staple food crops in the world. It has a wide range of cultivation, high yield, and high nutritional value. Enhancing the photosynthesis of potato is particularly important as it leads to an increase in the potato yield. The light-harvesting pigment-binding protein complex is very important for plant photosynthesis. We identified 12 Stlhcb gene family members from the potato variety "Atlantic" using transcriptome sequencing and bioinformatics. The proteins encoded by the Stlhcb gene family have between 3358 and 4852 atomic number, a relative molecular weight between 24060.16 and 34624.54 Da, and an isoelectric point between 4.99 and 8.65. The RT-qPCR results showed that the 12 Stlhcb genes were expressed in a tissue-specific and time-dependent fashion under low light. The relative expression of the Stlhcb genes in the leaves was significantly higher than that in the stems and roots, and the relative expression of these genes first increased and then decreased with the prolongation of light exposure time. The Stcp24 gene with the highest expression was cloned, and an expression vector was constructed. A subcellular localization analysis was performed in tobacco and an overexpression experiment was performed in potato using an Agrobacterium-mediated method. The subcellular localization analysis showed that the protein encoded by Stcp24 was located in chloroplasts as expected. Overexpression of Stcp24 in transgenic potato increased the yield of potatoes and the content of chlorophyll a and b; increased the net photosynthetic rate, transpiration rate, stomatal conductance, electron transport efficiency, and semi-saturated light intensity; and promoted photosynthesis and plant growth. This study provides a reference for the study of the function of the potato light-harvesting pigment-binding protein gene family. It lays a foundation for further study of the mechanism of the photosynthesis of potato, improvement of the light energy utilization of potato, and molecular breeding of potato.

Solanum tuberosum↗

Photosynthesis and sink activity of wasp-induced galls in Acacia pycnantha.

Although insect galls are widely known to influence source-sink relationships in plants, the relationship between photosynthesis and gall activity has not been extensively studied. In this study we used 14CO2, photosynthesis, and respiration measurements to examine the capacity of bud galls induced by the wasp Trichilogaster signiventris (Pteromalidae) as carbon sinks in Acacia pycnantha. Galls of this species develop either in vegetative or reproductive buds, depending on the availability of tissues at different times of the year, and effectively eliminate seed production by the plant. Photosynthetic rates in phyllodes subtending clusters of galls were greater than rates in control phyllodes, a result we attributed to photosynthesis compensating for increased carbon demand by the galls. Contrary to previous studies, we found that photosynthesis within galls contributed substantially to the carbon budgets of the galls, particularly in large, mature galls, which exhibited lower specific respiration rates allowing for a net carbon gain in the light. To determine the sink capacity and competitive potential of galls, we measured the proportion of specific radioactivity in galls originating from either vegetative or reproductive buds and found no difference between them. The proportion of the total amount of phyllode-derived 14C accumulated in both clustered and solitary galls was less than that in fruits. Galls and fruits were predominantly reliant on subtending rather than on distant phyllodes for photosynthate. Solitary galls that developed in vegetative buds constituted considerably stronger sinks than galls in clusters on inflorescences where there was competition between galls or fruits for resources from the subtending phyllode. Wasps developing in solitary vegetative galls were correspondingly significantly larger than those from clustered galls. We conclude that, in the absence of inflorescence buds during summer and fall, the ability of the wasps to cause gall formation in vegetative tissues tempers intraspecific competition and substantially increases the availability of plant resources for the development of wasps in such galls.

Acacia↗

Photosynthesis and kinetic characteristics of rubisco in Hibiscus cannabinus L.

Photosynthetic characteristics in kenaf (Hibiscus cannabinus L.), a C3 plant, were compared with Abelmoschus esculentus (L.) Moench, another member of Malvaceae. Kenaf leaves exhibited significantly higher rate of photosynthesis (40 mg CO2 dm(-2) hr(-1)) which was 24.6 mg dm(-2) hr(-1) in A. esculentus. Rate of photo and dark respiration was similar in both the species. Kenaf leaf photosynthesis had a higher optimum temperature (32 degrees C) than that of A. esculentus (26 degrees C). Photosynthesis in kenaf leaves required higher saturation irradiance (1,600 micromole m(-2) sec(-1)). There was a significant correlation between photosynthetic rate and biomass yield in these species. The primary product of photosynthesis after 5 seconds of 14C-assimilation was 3-PGA in both the species. The kinetic properties of RuBP carboxylase/oxygenase were determined in the leaf extracts. Higher carboxylase activities were recorded with kenaf leaf extracts (245 pmole mg chl(-1) hr(-1)). Km (CO2) for kenaf leaf carboxylase was significantly lower (7.8 microM) than A. esculentus (13.5 microM) and corresponding difference in Vmax values of carboxylase was recorded between the two species. The kinetic characteristics of oxygenase were similar in both the extracts. These results indicated the variation in carboxylase activity and its kinetic characteristics reflected a significant difference in CO2 assimilation in C3 plants.

Hibiscus↗

[A comparative study on photosynthesis and transpiration of Kelimeris integrifolia in summer and autumn on Songnen grasslands of China].

The diurnal changes in photosynthesis and transpiration of Kelimeris integrifolia in Summer and Autumn on Songnen grasslands were characterized as a two-peak curve, but with some differences between two seasons. It was shown that environmental conditions and plant internal factors had a strong effects on plant photosynthesis and transpiration. The net photosynthetic and transpiration rates of the leaves were positively correlated to photosynthetically active radiation (P < 0.01), and negatively correlated to stomatal resistance as well as intercellular CO2 concentration (P < 0.05). The transpiration rate was positively correlated to leaf temperature and water saturation deficit of the air. Among the above factors, photosynthesis active radiation was the dominant factor, and stomatal resistance also played an important role in plant photosynthesis and transpiration.

China↗

[Effects of different light transmission rate on American ginseng's photosynthesis].

From 1997 to 1998, series of experiments were carried out to study American ginseng's photosynthesis related to light transmission rate of plastic cover. The results showed that American ginseng's light saturation point for photosynthesis was different under the different light transmission rate (LTR) because of shading. At about 29.0 degrees C and under 12%, 30%, 42% LTR, 4-year-old American ginseng's light saturation point were 171.0, 323.0, and 429.0 mumol.m-2.s-1, respectively. The maximum net photosynthesis rate (Pn) was 6.54 mg CO2.dm-2.h-1, which appeared under the shading of 30% LTR. The daily course of 3-year-old American ginseng's Pn changed with LTR. When LTR was less than 25.8%, the daily curves of Pn had a single peak, and when LTR was higher than 25.8%, it had two peaks and the leaf's photosynthesis noon break phenomenon was remarkable. The results of single correlative analysis showed that the dominant factor that influenced Pn was photon flux density (PFD), and the results of plural regression analysis showed that the synthetic effect of all the influencing factors together on Pn was significant.

Light↗

Increased expression of a gene coding for NAD:glyceraldehyde-3-phosphate dehydrogenase during the transition from C3 photosynthesis to crassulacean acid metabolism in Mesembryanthemum crystallinum.

We utilized differential plaque hybridization to identify three cDNA clones for transcripts which increase in abundance during the salinity-induced transition from C3 photosynthesis to crassulacean acid metabolism (CAM) in Mesembryanthemum crystallinum. Although there are differences in the abundance of these transcripts in unstressed tissue, steady-state levels of all three increased within 30 h following irrigation with 0.5 M NaCl. One cDNA encodes the cytosolic form of glyceraldehyde-3-phosphate dehydrogenase (D-glyceraldehyde-3-phosphate:NAD+ oxidoreductase (phosphorylating) (NAD-GAPDH], an enzyme involved in the production of phosphoenolpyruvate for CO2 fixation at night and the conversion of pyruvate to storage carbohydrate during the day. Coding region and 3'-noncoding sequence probes were used to examine the expression of NAD-GAPDH transcripts in leaf and root tissue. We show that the gene encoding the NAD-GAPDH cDNA is expressed in both leaf and root tissue during C3 photosynthesis and CAM. NAD-GAPDH transcript levels increase rapidly in leaf (but not in root) tissue during the transition to CAM. Our data indicate that the predominant NAD-GAPDH transcript expressed during C3 photosynthesis and CAM is encoded by a single gene in M crystallinum. These results imply that the transition to CAM in some cases involves an upward readjustment in the level of a gene product expressed during C3 photosynthesis, rather than the expression of a CAM-specific isoform with unique regulatory or kinetic properties.

Amino Acid Sequence↗

Modeling effects of Photoadaption on the Photosynthesis-Irradiance Curve

In this article, I analyse how photoadaption affects the photosynthesis-irradiance curve in phytoplankton. Four parameters are presumably affected by photoadaption: (1) the size of a photosynthetic unit; (2) the number of photosynthetic units per cell; (3) the average turnover time of a photosynthetic unit; and (4) the chlorophyll-specific absorption cross-section. Prezelin's well-known conceptual model of photoadaption deals with variation in size and number of photosynthetic units, but not with the other two parameters. Prezelin's model predicts that the photosynthesis-irradiance curve is differentially affected by variations in size or number of photosynthetic units. By contrast, my analysis shows that if photoadaptional variation in the turnover time is also accounted for, only the cellular chlorophyll concentration is relevant; variation in the size vs. number of photosynthetic units is immaterial. The photoadaptional response of the chlorophyll-specific absorption cross-section allows a simple description consistent with experimental data. I use this description to derive a single expression for the rate of photosynthesis as dependent on irradiance and the cellular chlorophyll concentration. This model for the photosynthesis-irradiance-curve accounts for the photoadaptional alterations of all four parameters. Copyright 1997 Academic Press Limited

Journal Article↗

Influence of Algal Photosynthesis on Biofilm Bacterial Production and Associated Glucosidase and Xylosidase Activities.

Natural photosynthetic biofilms were incubated under light (100 mmol m-2 s-1) and dark conditions to elucidate the impact of photosynthesis on bacterial production, abundance, biovolume, biomass, and enzyme activities over 24 h. Use of organic carbon-free media limited carbon sources to algal photosynthesis and possibly the polysaccharides of the biofilm matrix. Bacterial production of biofilm communities was significantly higher in light incubations (p <0.001). The greatest differences in production rates between light and dark incubations occurred between 8 and 24 h. Biomass-specific a- and b-glucosidase and b-xylosidase activities were stimulated by photosynthesis, with significantly greater activities occurring at hours 16 and 24 in the light treatment (p <0.01). The results indicate that algal photosynthesis can have a significant impact on bacterial productivity, biomass, biovolume, and enzyme production over longer time periods at low photon flux densities (?100 mmol m-2 s-1).

Journal Article↗

Reduction of net photosynthesis in oats after treatment with low concentrations of ozone.

Oats (Avena sativa L. cv Titus) were exposed to low concentrations of O3 in an assimilation chamber system. Net photosynthesis (net CO2 uptake), measured before and after O3 fumigation, showed significantly different responses for leaves of different age. The oldest active leaf was the most sensitive to O3. Net photosynthesis was depressed after 2 h with 0.075 ppm (150 microg m(-3)) O3. For leaves exposed to 0.150 ppm (300 microg m(-3)) O3 for 2 h, net photosynthesis was reduced significantly for 4 h, after which recovery occurred, nearly reaching the preexposure level 19 h after the exposure. Dark respiration was initially more than doubled after exposure to 0.130 ppm (260 microg m(-3)) O3. There was no visible injury after any of the experiments. The results indicate that O3 may cause crop losses through effects on photosynthesis even in Scandinavia, where a typical O3 episode lasts 1 to 2 h, and the concentration seldom exceeds 0.150 ppm.

Journal Article↗

Photosynthesis is limited at high leaf to air vapor pressure deficit in a mutant of Arabidopsis thaliana that lacks trienoic fatty acids.

We have investigated the role of polyunsaturated fatty acids in photosynthesis using a triple mutant of Arabidopsis thaliana that lacks trienoic fatty acids (fad 3-2 fad 7-2 fad 8). Though this mutant is male sterile, vegetative growth and development under normal conditions are largely unaffected (McConn and Browse, 1996 Plant Cell 8: 403-416). At 0.2-1.0 kPa vapor pressure deficit (low VPD), maximum photosynthetic rates of wild-type and mutant plants were similar while stomatal conductance rates were up to 2 times higher in mutant plants. However, light-saturated rates of carbon assimilation and stomatal conductance in the mutant were lower than in wild-type plants when measured at ambient (35 Pa) CO(2) and 2.0-2.8 kPa vapor pressure deficit (high VPD). The limitation to photosynthesis in the mutant plants at high VPD was overcome by saturating partial pressures of CO(2) suggesting a stomatal limitation. Chlorophyll fluorescence measurements indicate that differences observed in maximum assimilation rates were not due to limitations within the photochemical reactions of photosynthesis. Stomatal response to VPD and intrinsic water use efficiency was drastically different in mutant versus wild-type plants. The results of this investigation indicate that for Arabidopsis, polyunsaturated fatty acids may be an important determinant of responses of photosynthesis and stomatal conductance to environmental stresses such as high VPD.

Journal Article↗

Celebrating the millennium - historical highlights of photosynthesis research, Part 2.

This paper is an introduction to Part 2 of our celebrations of the historical highlights of photosynthesis research. Part 1 was published in October 2002 as Volume 73 of Photosynthesis Research. After a brief introduction, we recognize two giants in the field: Cornelis B. van Niel (for anoxygenic photosynthesis), and Robert Hill (for oxygenic photosynthesis). This is followed by recognition of a 1960 book by Hans Gaffron, and a multi-authored book edited by W. Ruhland and André Pirson, and inclusion in the appendix of a list of selected books. Our celebration is enhanced by the inclusion of beautiful paintings of cells by Antoinette Ryter. After introducing all the historical papers contained in this volume, we honor Louis N. M. Duysens, one of the greatest biophysicists of our time, and finally we dedicate this volume to a great scientist, humanist and peacemaker: Eugene I. Rabinowitch. [12pt] 'Annihilating all that is made To a green thought in a green shade' - Andrew Marvell (1621-1678), The Garden (1681).

Journal Article↗

Celebrating the millennium: historical highlights of photosynthesis research, part 3.

This paper introduces the third and final part of the 'millennium celebrations of historical highlights of photosynthesis research.' Part 1 (308 pages) was published in October 2002 as Vol. 73 of the journal Photosynthesis Research, and Part 2 (458 pages) was published in July 2003 as Vol. 76. Here, we recognize particularly the work of three major contributors to our understanding of photosynthesis: Roger Stanier (1916-1982); Germaine Cohen-Bazire (Stanier) (1920-2001); and William Arnold (1904-2001). We also introduce the historical papers contained in this volume; consider the legacy of Alfred Nobel (1833-1896); and identify Nobel prizes of special relevance to understanding the capture, conversion, and storage of light energy in both anoxygenic and oxygenic photosynthesis.

Journal Article↗

Influence of CO2 and sucrose on photosynthesis and transpiration of Actinidia deliciosa explants cultured in vitro.

Explants of Actinidia deliciosa Chev. Liang and Ferguson var. Hayward were cultured in controlled CO2 atmospheres in the presence of different sucrose concentrations. Organogenesis was measured after 45 days in explants from the different assays, and quantification of photosynthesis, transpiration, chlorophylls, RUBISCO and total soluble protein content was performed in leaves from the different treatments. The best results were those of explants cultured at 600 &mgr;mol CO2 mol-1 on 20 g l-1 of sucrose for the first 20 days and then transferred to sucrose-free medium until the end of the culture period. Increasing CO2 to 2000 &mgr;mol CO2 mol-1 in the atmosphere of the culture vessel reduced all the parameters studied. Photosynthesis of autotrophically developed explants trebled that of the reference heterotrophic explants, as there was an apparent inverse relationship between photosynthesis and transpiration. Photosynthesis was saturated at 300 &mgr;mol m-2 s-1 PPFD and 600 &mgr;mol CO2 mol-1. Chlorophylls and RUBISCO presented differences between treatments, mainly between different CO2 concentrations, with the highest values in autotrophically cultured explants. Explants grown at 2000 &mgr;mol CO2 mol-1 showed the lowest RUBISCO/Prots ratio, probably due to negative adaptation of RUBISCO to long-term high CO2. In short, explants grown in a controlled microenvironment, with increased CO2 and under autotrophic conditions, developed wholly functional photosynthetic apparatus well prepared to be transferred to ex vitro conditions, which has many advantages in micropropagation.

Journal Article↗

Changes in photosynthesis and water status of developing leaves of Brachystegia spiciformis Benth.

Changes in net carbon assimilation and water status were studied during leaf development in the deciduous, tropical species Brachystegia spiciformis Benth. In this upland savanna African tree, bud-burst and leaf development occur approximately two months before the rainy season. The newly formed leaves synthesize anthocyanin until the fully expanded leaves of the whole canopy are red. This foliage is referred to as "spring flush" foliage. Subsequently, the anthocyanins are metabolized and the pre-rain leaves become green. Carbon dioxide assimilation exhibited a bimodal diurnal pattern and was similar for pre-rain green leaves and fully expanded flushing leaves, although pre-rain green leaves showed a net uptake of carbon throughout the daylight period, whereas flushing leaves exhibited only brief periods of net photosynthesis in the morning and early afternoon. Measurements of leaf water potential and relative water content showed a diurnal pattern with considerable variation throughout the day. Leaf water potential and relative water content values decreased soon after sunrise reaching a minimum at a time corresponding to the afternoon peak in CO(2) assimilation. Stomatal conductance was closely related to transpiration rate in both flushing and pre-rain green leaves, although flushing leaves had lower stomatal conductances than pre-rain green leaves. Pre-rain green leaves exhibited a compensation irradiance of approximately 180 micro mol m(-2) s(-1), whereas flushing leaves had positive net photosynthesis only at PPFDs greater than 300 micro mol m(-2) s(-1). Rate of photosynthesis (expressed per leaf area or chlorophyll unit) increased as anthocyanin concentration decreased, although the photosynthetic rate continued to increase long after the leaf anthocyanins had been degraded to low, visually undetectable amounts. Post-rain green leaves had chlorophyll concentrations, transpiration rates and stomatal conductances similar to those of pre-rain green leaves; however, photosynthetic rates in post-rain leaves were more than three times higher. Thus, during the early stages of the spring flush, carbon asimilation rates of the flushing leaves were inversely related to leaf anthocyanin concentrations. In pre-rain green leaves, photosynthesis was limited by other non-stomatal factors.

Journal Article↗

Parameterization and testing of a coupled photosynthesis-stomatal conductance model for boreal trees.

A coupled photosynthesis-stomatal conductance model was parameterized and tested with branches of black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.) trees growing in the Northern Study Area of the Boreal Ecosystem-Atmosphere Study (BOREAS) in Manitoba, Canada. Branch samples containing foliage of all age-classes were harvested from a lowland old black spruce (OBS) and an old jack pine (OJP) stand and the responses of photosynthesis (A(n)) and stomatal conductance (g(s)) to temperature, CO(2), light, and leaf-to-air vapor pressure difference (VPD) were determined under controlled laboratory conditions at the beginning, middle, and end of the growing season (Intensive Field Campaigns (IFC) 1, 2, and 3, respectively). The parameterized model was then tested against in situ field gas-exchange measurements in a young jack pine (YJP) and an upland black spruce (UBS) stand as well as in the OBS and OJP stands. Parameterization showed that Rubisco capacity (V(max)), apparent quantum yield (alpha') and Q(10) for sink limitation were the most crucial parameters for the photosynthesis sub-model and that V(max) varied among different measurement series in the laboratory. Verification of the model against the data used to parameterize it yielded correlation coefficients (r) of 0.97 and 0.93 for black spruce and jack pine, respectively, when IFC-specific parameters were used, and 0.77 and 0.87 when IFC-2 parameters were applied to all IFCs. For both measured and modeled g(s), the stomatal conductance sub-model, which linearly relates g(s) to (A(n)h(s))/c(s) (where h(s) and c(s) are relative humidity and CO(2) mole fraction at the leaf surface, respectively), had significantly steeper slopes and higher r values when only the VPD response data were used for parameterization than when all of the response data were used for parameterization. Testing the photosynthesis sub-model against upper canopy field data yielded poor results when laboratory estimates of V(max) were used. Use of the mean V(max) estimated for all upper canopy branches measured on a given day improved model performance for jack pine (from a nonsignificant correlation between measured and modeled A(n) to r = 0.45), but not for black spruce (r = 0.45 for both cases). However, when V(max) was estimated for each branch sample individually, the model accurately predicted the 23 to 137% diurnal variation in A(n) for all stands for both the upper and lower canopy. This was true both when all of the other parameters were IFC-specific (r = 0.93 and 0.92 for black spruce and jack pine, respectively) and when only mid-growing season (IFC-2) values were used (r = 0.92 for both species). Branch-specific V(max) estimates also permitted accurate prediction of field g(s) (r = 0.75 and 0.89 for black spruce and jack pine, respectively), although parameterization with all of the response data overestimated g(s) in the field, whereas parameterization with only the VPD response data provided unbiased predictions. Thus, after parameterization with the laboratory data, accurately modeling the range of A(n) and g(s) encountered in the field for both black spruce and jack pine was reduced to a single unknown parameter, V(max).

Journal Article↗