Vacuolar pH oscillations in mesophyll cells accompany oscillations of photosynthesis in leaves: Interdependence of cellular compartments,and regulation of electron flow in photosynthesis

Oscillations of photosynthesis induced in leaves of Vicia faba L. were accompanied by oscillations not only in the pH of the chloroplast stroma, but also by pH oscillations in the cytosol and in the vacuole of leaf mesophyll cells. Cytosolic pH oscillations were in phase with stromal oscillations, but antiparallel to vacuolar pH oscillations. During maxima of photosynthesis, the cytosolic pH exhibited maxima and the vacuolar pH minima. Vacuolar acidification is interpreted to be the result of energized proton transport from the cytosol into the vacuole. Since the ratio of dihydroxyacetone phosphate to phosphoglycerate is maximal during the peaks of photosynthesis (Stitt et al., 1988, J. Plant Physiol. 133, 133–143; Laisk et al., 1991, Planta 185, 554–562), while the activity of NADP-malic dehydrogenase is highest during minima of photosynthesis (Scheibe and Stitt, 1988, Plant Physiol. Biochem. 26, 473–481), the present data indicate in agreement with earlier observations (Yin et al., 1991, Planta 184, 30–34) that light-dependent cytosolic energization is brought about by the oxidation of dihydroxyacetone phosphate rather than of malate. They also indicate that the over-reduction of the electrontransport chain observed during minima of photosynthesis is relieved not predominantly by oxaloacetate reduction and export of the resulting malate from the chloroplasts but by another reaction, presumably oxygen reduction.Abbreviations CDCF 5-(and 6-)carboxy-2,7-dichlorofluorescein     

Light-harvesting properties of zinc complex of chlorophyll-<Emphasis Type="Italic">a</Emphasis> from <Emphasis Type="Italic">spirulina</Emphasis> in surfactant micellar media

Zn chlorophyll-a was prepareted from Mg chlorophyll-a from spirulina and the optical properties of the ground state and the photoexited state of Zn chlorophyll-a in aqueous surfactant micellar media were studied using UV-vis absorption, fluorescence emission spectra, electrochemical and fluorescence lifetime measurements. In comparison of the UV-vis absorption and fluorescence emission spectra of Zn chlorophyll-a and Mg chlorophyll-a, the blue-shift in the absorption bands and emission peak of Zn chlorophyll-a was observed. The energies of the first excited singlet state of Zn chlorophyll-a was 1.87eV. The first oxidation and reduction potentials of the photoexcited singlet state of Zn chlorophyll-a were −0.67 and 0.60V, respectively. Fluorescence lifetime of Zn chlorophyll-a was 9.0 ns in CTAB micellar solution. The fluorescence lifetime of Zn chlorophyll-a is shorter than that of Mg chlorophyll-a (9.8 ns). The photositability of Zn chlorophyll-a was superior to that of Mg chlorophyll-a in various pH conditions. Published online December 2004     

Primary productivity and related parameters in three different types of inland waters in Sri Lanka

Gross and net primary production together with chlorophyll-a biomass were investigated with respect to depth and diurnal changes in three categories of inland waters (reservoirs, temporary ponds, brackish water lagoons) in Sri Lanka. Ten field sites, in both the dry and wet zones of the island, were investigated. Bimodal productivity profiles were recorded in two of the three reservoirs studied. The diel pattern of net photosynthetic rate varied between sites although peak photosynthetic efficiency occurred at solar noon. Surface photoinhibition was characteristic of the reservoirs and brackish water lagoons but not of the temporary ponds. Mean gross primary production was 3.02 g C m^–2 d^–1 but was higher in the temporary ponds than in the reservoirs. The gross primary production in the brackish water Koggala Lagoon at 0.08 g C m^–2 d^–1 is a record low for tropical lagoons and was 2.5 times less than the two other lagoons investigated. Variability in net primary production between sites was similar to the variation in gross production with a relatively low mean value for tropical inland waters of 0.495 C m^–2 d^–1. Mean maximum photosynthetic rate was 0.30 mg C m^–3 h^–1 but was lower in the reservoirs than in the temporary ponds and lagoons.     

Pollen limitation and distance-dependent fecundity in females of the clonal gynodioecious herb Glechoma hederacea (Lamiaceae)

Summary Pollen movement is often restricted in natural populations, and insufficient pollination is a potential constraint on sexual reproduction in outcrossing species. Seed-set should decrease with increased distance from the pollen source in outcrossing plants. This prediction was tested using females of the clonal, gynodioecious herb Glechoma hederacea in three natural populations. In controlled pollinations, both hermaphrodites and females had similar high percentages of fruit-set and seed-set. In a natural population where a female clone was isolated from the nearest hermaphroditic clone by c. 100 m, fruit-set was low (1%). In another population where hemaphroditic clones were rare and female clones had a patchy distribution, fruit-and seed-set in females were pollen-limited and decreased with increased distance from the nearest pollen source. The estimated mean pollen dispersal distance was 5.9 m when calculated on fruit-set and 5.3 m when calculated on seed-set. The most frequent pollinators were bumblebees. The mean and median distances moved by pollinators between ramets were 0.13 m and 0.05 m. In a third population where female clones were isolated from the nearest hermaphrodites by more than 200 m, fruit-set was 0%. After introduction of 16 hermaphroditic ramets in the center of the female clone, fruit-set varied between 0% and 100% in individual female ramets. Fruit-set decreased with increased distance from the pollen source. The mean and median pollen movement distances were 1.06 m and 0.54 m.     

Environmental heterogeneity and clonal growth: a study of the capacity for reciprocal translocation in Glechoma hederacea L.

Clonal fragments of Glechoma hederacea L. (Lamiaceae) were subjected to environments in which light and nutrients were supplied with a strictly negative association in space, i.e. when one of these resources was in ample supply the other was scarce. Treatments were chosen to simulate environments in which clones grew either within homogeneous conditions or across patch types (heterogeneous conditions). The hypothesis was tested that reciprocal translocation (i.e. exchange of both nutrients and assimilates) between connected groups of ramets would increase biomass production of clones growing under heterogeneous conditions compared to that of clones growing in homogeneous conditions. A cost-benefit analysis was carried out to test this hypothesis. Results suggested that reciprocal translocation did not occur at the structural scale considered in this experiment; no evidence was found for a significant effect on whole clone biomass of assimilate and/or nutrient translocation between clone parts experiencing contrasting levels of resource supply. It is suggested that predominantly acropetal movement of resources and the pattern of integrated physiological unit formation in G. hederacea are the main properties responsible for the lack of mutual physiological support between connected clonal fragments growing in differing habitat conditions. These properties are expected to promote clonal expansion and the exploitation of new territory, rather than sustaining clone parts in sub-optimal patches of habitat for prolonged periods of time.     

Phosphate sequestration by glycerol and its effects on photosynthetic carbon assimilation by leaves

Glycerol induced a limitation on photosynthetic carbon assimilation by phosphate when supplied to leaves of barley (Hordeum vulgare L.) and spinach (Spinacia oleracea L.). This limitation by phosphate was evidenced by (i) reversibility of the inhibition of photosynthesis by glycerol by feeding orthophosphate (ii) a decrease in light-saturated rates of photosynthesis and saturation at a lower irradiance, (iii) the promotion of oscillations in photosynthetic CO₂ assimilation and in chlorophyll fluorescence, (iv) decreases in the pools of hexose monophosphates and triose phosphates and increases in the ratio of glycerate-3-phosphate to triose phosphate, (v) decreased photochemical quenching of chlorophyll fluorescence, and increased non-photochemical quenching, specifically of the component which relaxed rapidly, indicating that thylakoid energisation had increased. In barley there was a massive accumulation of glycerol-3-phosphate and an increase in the period of the oscillations, but in spinach the accumulation of glycerol-3-phosphate was comparatively slight. The mechanism(s) by which glycerol feeding affects photosynthetic carbon assimilation are discussed in the light of these results.Abbreviations Chl chlorophyll - C _i intercellular concentration of CO₂ - P phosphate - PGA glycerate-3-phosphate - Pi orthophosphate - triose-P sum of glyceraldehyde-3-phosphate and dihydroxyacetone phosphate     

Effects of temperature on the regulation of photosynthetic carbon assimilation in leaves of maize and barley

The aim of this work was to examine the effect of temperature in the range 5 to 30 ° C upon the regulation of photosynthetic carbon assimilation in leaves of the C₄ plant maize (Zea mays L.) and the C₃ plant barley (Hordeum vulgare L.). Measurements of the CO₂-assimilation rate in relation to the temperature were made at high (735 bar) and low (143 bar) intercellular CO₂ pressure in barley and in air in maize. The results show that, as the temperature was decreased, (i) in barley, pools of phosphorylated metabolites, particularly hexose-phosphate, ribulose 1,5-bisphosphate and fructose 1,6-bisphosphate, increased in high and low CO₂; (ii) in maize, pools of glycerate 3-phosphate, triose-phosphate, pyruvate and phosphoenolpyruvate decreased, reflecting their role in, and dependence on, intercellular transport processes, while pools of hexose-phosphate, ribulose 1,5-bis phosphate and fructose 1,6-bisphosphate remained approximately constant; (iii) the redox state of the primary electron acceptor of photosystem II (Q_A) increased slightly in barley, but rose abruptly below 12° C in maize. Non-photochemical quenching of chlorophyll fluorescence increased slightly in barley and increased to high values below 20 ° C in maize. The data from barley are consistent with the development of a limitation by phosphate status at low temperatures in high CO₂, and indicate an increasing regulatory importance for regeneration of ribulose 1,5-bisphosphate within the Calvin cycle at low temperatures in low CO₂. The data from maize do not show that any steps of the C₄ cycle are particularly cold-sensitive, but do indicate that a restriction in electron transport occurs at low temperature. In both plants the data indicate that regulation of product synthesis results in the maintenance of pools of Calvin-cycle intermediates at low temperatures.Abbreviations Glc6P glucose-6-phosphate - Fru6P fructase-6-phosphate - Frul,6bisP fructose-1,6-bisphosphate - PGA glycerate-3-phosphate - p _i intercellular partial pressure of CO₂ - RuBP ribulose-1,5-bisphosphate - triose-P sum of glyceraldehyde-3-phosphate and dihydroxyacetone phosphate We thank the Agricultural and Food Research Council, UK (Research grant PG50/67) and the Science and Engineering Research Council, UK for financial support. C.A.L. was supported by the British Council, by the Conselho Nacional de Desenvolvimento Cientiflco e Tecnologico (CNPq), Brazil and by an Overseas Research Student Award. We also thank Mark Stitt (Bayreuth, FRG) and Debbie Rees for helpful discussions.     

An analysis of the costs and benefits of physiological integration between ramets in the clonal perennial herb Glechoma hederacea

Summary The costs and benefits, measured in terms of dry weight, of physiological integration between clonal ramets, were analysed in two experiments conducted on the clonal herb Glechoma hederacea. Firstly, integration between consecutively-produced ramets was examined in an experiment in which stolons grew from one set of growing conditions (either unshaded or shaded and either nutrient-rich or nutrient-poor) into conditions in which light or nutrient level was altered. Comparisons were made between the dry weight of the parts of the clones produced before and after growing conditions were changed, and the dry weights of the corresponding part of control clones subjected to constant growing conditions. In a second experiment, integration between two distinct parts of G. hederacea clones was investigated. In this experiment clones were grown from two connected parent ramets and the parts of the clone produced by each parent ramet were subjected independently to either nutrient-rich or nutrient-poor conditions. Ramets in resource-rich conditions provided considerable physiological support to those in resource-poor conditions. This was measured as a dry weight gain compared with the weight of the corresponding part of the control clones growing in resource-poor conditions. However, when stolons grew from resource-poor conditions into resource-rich conditions, there was no similar evidence of the resourcepoor ramtes receiving support from resource-rich ramets. Physiological integration did not result in dry weight gains when this would have necessitated basipetal translocation of resources.Because of the predominantly acropedal direction of movement of translocates in G. hederacea, the structure of the clone was important in determining the effectiveness of integration between ramets. Where physiological integration was effective, the cost to the supporting ramets in terms of dry weight was insignificant. Physiological integration allows clones to maintain a presence in less favourable sites with insignificant cost to ramets in favourable sites, thereby reducing the probability of invasion by other plants, and providing the potential for rapid clonal growth if conditions improve. Integrated support of ramets in unfavourable conditions also enables the clone to grow through unfavourable sites, thus increasing the probability of encountering more favourable conditions by wider foraging.     

Phytoplankton productivity and chlorophyll-A concentration of Oguta Lake in Southeastern Nigeria

The phytoplankton productivity and chlorphyll-a concentration of Oguta Lake, the largest natural lake in south-eastern Nigeria, are presented (Dec. 1983. Nov. 1984). The gross productivity ranged from 1.3 to 3.77 g C.m^–2.day^–1 for the water column, dropped during the period of heavy rainfall and varied with depth. The chlorophyll-a concentration had monthly means ranging from 2.31 to 4.00 mg.m^–3, with a drop during the rains, but little depth variation. Both productivity and chlorophyll-a showed non-significant correlation with the physico-chemical features of the water. The values of the biological parameters showed the lake as mesotrophic. The values are compared with those of other African lakes.     

The relationship between contents of photosynthetic metabolites and the rate of photosynthetic carbon assimilation in leaves of Amaranthus edulis L.

The relationship between the gas-exchange characteristics of attached leaves of Amaranthus edulis L. and the contents of photosynthetic intermediates was examined in response to changing irradiance and intercellular partial pressure of CO₂. After determination of the rate of CO₂ assimilation at known intercellular CO₂ pressure and irradiance, the leaf was freeze-clamped and the contents of ribulose-1,5-bisphosphate, glycerate-3-phosphate, fructose-1,6-bisphosphate, glucose-6-phosphate, fructose-6-phosphate, triose phosphates, phosphoenolpyruvate, pyruvate, oxaloacetate, aspartate, alanine, malate and glutamate were measured. A comparison between the sizes of metabolite pools and theoretical calculations of metabolite gradients required for transport between the mesophyll and the bundle-sheath cells showed that aspartate, alanine, glycerate-3-phosphate and triose phosphates were present in sufficient quantities to support transport by diffusion, whereas pyruvate and oxaloacetate were not likely to contribute appreciably to the flux of carbon between the two cell types. The amounts of ribulose-1,5-bisphosphate were high at low intercellular partial pressures of CO₂, and fell rapidly as the CO₂-assimilation rate increased with increasing intercellular partial pressures of CO₂, indicating that bundle-sheath CO₂ concentrations fell at low intercellular partial pressures of CO₂. In contrast, the amount of phosphoenolpyruvate and of C₄-cycle intermediates declined at low intercellular partial pressures of CO₂. This behaviour is discussed in relation to the co-ordination of carbon assimilation between the Calvin and C₄ cycles.Abbreviations PEP phosphoenolpyruvate - PGA glycerate-3-phosphate - p _i intercellular CO₂ pressure - RuBP ribulose-1,5-bisphosphate - triose-P triose phosphates     

Chlorophyll fluorescence imaging of photosynthetic activity in sun and shade leaves of trees

The differences in pigment levels, photosynthetic activity and the chlorophyll fluorescence decrease ratio R _Fd (as indicator of photosynthetic rates) of green sun and shade leaves of three broadleaf trees (Platanus acerifolia Willd., Populus alba L., Tilia cordata Mill.) were compared. Sun leaves were characterized by higher levels of total chlorophylls a + b and total carotenoids x + c as well as higher values for the weight ratio chlorophyll (Chl) a/b (sun leaves 3.23–3.45; shade leaves: 2.74–2.81), and lower values for the ratio chlorophylls to carotenoids (a + b)/(x + c) (with 4.44–4.70 in sun leaves and 5.04–5.72 in shade leaves). Sun leaves exhibited higher photosynthetic rates P _N on a leaf area basis (mean of 9.1–10.1 μmol CO₂ m⁻² s⁻¹) and Chl basis, which correlated well with the higher values of stomatal conductance G _s (range 105–180 mmol m⁻² s⁻¹), as compared to shade leaves (G _s range 25–77 mmol m⁻² s⁻¹; P _N: 3.2–3.7 μmol CO₂ m⁻² s⁻¹). The higher photosynthetic rates could also be detected via imaging the Chl fluorescence decrease ratio R _Fd, which possessed higher values in sun leaves (2.8–3.0) as compared to shade leaves (1.4–1.8). In addition, via R _Fd images it was shown that the photosynthetic activity of the leaves of all trees exhibits a large heterogeneity across the leaf area, and in general to a higher extent in sun leaves than in shade leaves.     

Analysis of the deep chlorophyll maximum across the Azores Front

Physical, chemical and biological observations made in late July and August 1997 across the Azores Front (37° N, 32°W to 32° N, 29°W) are presented. The objectives of the study were: (1) to analyse horizontal and vertical profiles of temperature, salinity, density, nutrients and chlorophyll-a (Chl a) of the top 350 m; (2) to identify the main differences in the deep Chl a Maximum (DCM) and hydrographic structure between the water masses that pass north and south of the Azores Front; and (3) to estimate phytoplankton primary production in these water masses. Horizontal and vertical profiles of salinity, temperature, density, nutrients and phytoplankton pigments in the top 350 m were analysed. The Front separates two distinct water types: the 18°C Mode Water (18 MW) of sub-tropical origin, and the 15°C Mode Water (15 MW) of sub-polar origin. Differences in the DCM and hydrographic structure between 18 MW and 15 MW were observed in the contour plots of each section. The average Chl a concentration between 5 and 200 m depth decreased significantly from 15 MW to 18 MW. The same pattern was observed for the Chl a concentration at the DCM depth. A vertical one-dimensional model was used to estimate the phytoplankton primary production in the 15 MW and 18 MW and led to an estimated water column average gross primary productivity (GPP) between 1.08 and 2.71 mg C m^–3 d^–1 for the 15 MW and about half of these values for the 18 MW. These results indicate that the typical south–north positive slope on DCM depth parallels a latitudinal increase on GPP, suggesting that the location of the Azores Front may have a significant regional impact on GPP.     

Analysis of oxygen evolution during photosynthetic induction and in multiple-turnover flashes in sunflower leaves

Exchange of CO₂ and O₂ and chlorophyll fluorescence were measured in the presence of 360 1 · 1^–1 CO₂ in nitrogen in Helianthus annuss L. leaves which had been preconditioned in the dark or at a photon flux density (PFD) of 24 mol · m^–2 · s^–1 either in 21 or 0% O₂. An initial light-dependent O₂ outburst of 6 mol · m^–2 was measured after aerobic dark incubation. It was attributed to the reduction of electron carriers, predominantly plastoquinone. The maximum initial rate of O₂ evolution at PFD 8000 mol · m^–2 · s^–1 was 170 mol · m^–2 · s^–2 or about four times the steady CO₂-and light-saturated rate of photosynthesis. Fluorescence measurements showed that the rate was still acceptor-limited. Fast O₂ evolution ceased after electron carriers were reduced in the dark-adapted leaf, but continued for a short time at the lower rate of 62 mol · m^–2 · s^–1 in the light-adapted leaf. The data are interpreted to show that enzymes involved in 3-phosphoglycerate reduction are dark-inhibited, but were fully active in low light. In a dark-adapted leaf, respiratory CO₂ evolution continued under nitrogen; it was partially inhibited by illumination. Prolonged exposure of a leaf to anaerobic conditions caused reducing equivalents to accumulate. This was shown by a slowly increasing chlorophyll fluorescence yield which indicated the reduction of the PSII acceptor Q_A in the dark. When the leaf was illuminated, no O₂ evolution was detected from short light pulses, although transient O₂ production was appreciable during longer light pulses. This indicates that an electron donor (pool size about 2–3 e/PSII reaction center) became reduced in the dark and the first photons were used to oxidise this donor instead of water.Abbreviations Chl chlorophyll - CRC carbon reduction cycle - GAPDH NADP-glyceraldehyde-phosphate dehydrogenase - PFD photon flux density - PGA 3-phosphoglycerate - RuBP ribulose bisphosphate - TCA tricarboxylic acid cycle To whom correspondence should be addressedThis work received support by the Estonian Academy of Sciences, the Gottfried-Wilhelm-Leibniz Program of the Deutsche For-schungsgemeinschaft and the Sonderforschungsbereich 251 of the University of Würzburg.     

The photosynthetic induction response in wheat leaves: net CO2 uptake,enzyme activation,and leaf metabolites

The photosynthetic induction response was studied in whole leaves of wheat (Triticum aestivum L.) following 5-min, 30-min and 10-h dark periods. After the 5-min dark treatment there was a rapid burst in the rate of photosynthesis upon illumination (half of maximum after 30s), followed by a slight decrease after 1.5 more min and then a gradual rise to the maximum rate. During this initial burst in photosynthesis, there was a rapid rise in the level of 3-phosphoglycerate (PGA) and a high PGA/triose-phosphate (triose-P) ratio was obtained. In addition, after the 5-min dark treatment, ribulose-1,5-bisphosphate carboxylase (Rubisco, EC 4.1.1.39), ribulose-5-phosphate kinase (EC 2.7.1.19) and chloroplastic fructose-1,6-bisphosphatase (EC 3.1.3.11) maintained a relatively high state of activation, and maximum activation occurred within 1 min of illumination. The results indicate there is a high capacity for CO₂ fixation in the cycle upon illumination but attaining maximum rates requires an increase in the ribulose-1,5-bisphosphate (RuBP) pool (adjustment in triose-P utilization for carbohydrate synthesis versus RuBP synthesis). With both the 30-min and 10-h dark pretreatments there was only a slight rise in photosynthesis upon illumination, followed by a lag, then a gradual increase to steady-state (half-maximum rate after 6 min). In contrast to the 5-min dark treatment, the level of PGA was low and actually decreased initially, whereas the level of RuBP increased and was high during induction, indicating that Rubisco is limiting. This regulation via the carboxylase was not reflected in the initial extractable activity, which reached a maximum by 1 min after illumination. The light activation of chloroplastic fructose-1,6-bisphosphatase in leaves darkened for 30 min and 10 h prior to illumination was relatively slow (reaching a maximum after 8 min). However, this was not considered to limit carbon flux through the carbon-fixation cycle during induction since RuBP was not limiting. When photosynthesis approached the maximum steady-state rate, a high PGA/triose-P ratio and a high PGA/RuBP ratio were obtained. This may allow a high rate of photosynthesis by producing a favorable mass-action ratio for the reductive phase (the conversion of PGA to triose phosphate) while stimulating starch and sucrose synthesis.Abbreviations Chl chlorophyll - FBP fructose-1,6-bisphosphate - FBPase fructose-1,6-bisphosphatase - Fru6P fructose-6-phosphate - Glc6P glucose-6-phosphate - PGA 3-phosphoglycerate - Pi inoganic phosphate - Rubisco RuBP carboxylase/oxygenase - RuBP ribulose-1,5-bisphosphate - Ru5P ribulose-5-phosphate - triose-P triose phosphates (dihydroxyacetone phosphate+glyceraldehyde-3-phosphate)     

Simultaneous and independent effects of abscisic acid on stomata and the photosynthetic apparatus in whole leaves

(±)-Abscisic acid (ABA) at 10^-5 M was added to the transpiration stream of leaves of 16 species (C₃ and C₄, monocotyledons and dicotyledons). Stomatal responses followed one of three patterns: i) stomata that were wide and insensitive to CO₂ initially, closed partially and became sensitive to CO₂; ii) for stomata that were sensitive to CO₂ before the application of ABA, the range of highest sensitivity to CO₂ shifted from high to low intercellular partial pressures of CO₂, for instance in leaves of Zea mays from 170–350 to 70–140 bar; iii) when stomata responded strongly to ABA, their conductance was reduced to a small fraction of the initial conductance, and sensitivity to CO₂ was lost. The photosynthetic apparatus was affected by applications of ABA to various degrees, from no response at all (in agreement with several previous reports on the absence of effects of ABA on photosynthesis) through a temporary decrease of its activity to a lasting reduction. Saturation curves of photosynthesis with respect to the partial pressure of CO₂ in the intercellular spaces indicated that application of ABA could produce three phenomena: i) a reduction of the initial slope of the saturation curve (which indicates a diminished carboxylation efficiency); ii) a reduction of the level of the CO₂-saturated rate of assimilation (which indicates a reduction of the ribulose-1,5-bisphosphate regeneration capacity); and iii) an increase of the CO₂ compensation point. Photosynthesis of isolated mesophyll cells was not affected by ABA treatments. Responses of the stomatal and photosynthetic apparatus were usually synchronous and often proportional to each other, with the result that the partial pressure of CO₂ in the intercellular spaces frequently remained constant in spite of large changes in conductance and assimilation rate. Guard cells and the photosynthetic apparatus were able to recover from effects of ABA applications while the ABA supply continued. Recovery was usually partial, in the case of the photosynthetic apparatus occasionally complete. Abscisic acid did not cause stomatal closure or decreases in the rate of photosynthesis when it was applied during a phase of stomatal opening and induction of photosynthesis that followed a transition from darkness to light.Abbreviations and symbols A rate of CO₂ assimilation - ABA (±)-abscisic acid - c _a partial pressure of CO₂ in the ambient air or in the gas supplied to the leaf chambers - c _i partial pressure of CO₂ in the intercellular spaces of a leaf - e _a partial pressure of H₂O in the air - g conductance for water vapor - J quantum flux - T ₁ leaf temperature     

Mutual shading and the photosynthetic capacity of exposed leaves of field grown soybeans

Light-saturated photosynthetic rates at air levels of carbon dioxide were measured about weekly in upper canopy leaves of two soybean cultivars grown at stand densities of 40 and 100 plants per square meter. Early in the season, when leaf area indices differed between stand densities, plants of both cultivars grown at high stand density had photosynthetic rates which averaged 23% lower than plants at low stand density. Later in the season, when there were no differences in leaf area index between stand densities, there were no differences in photosynthetic rates in the cultivar Kent, but rate differences of about 14% persisted in the cultivar Williams. In Williams mainstem leaves emerged into full sunlight later in their development at high than at low stand density. In both cultivars the oldest fully exposed leaves were photosynthetically immature for much of the season, as higher rates could be achieved by lower leaves which were shaded in situ. The results identify shading of young developing leaves and photosynthetic immaturity of fully exposed leaves as factors limiting canopy photosynthesis in soybeans, and indicate cultivar differences in how much high stand density reduces photosynthetic capacity.     

Controls on chlorophyll-a in nutrient-rich agricultural streams in Illinois, USA

Nitrogen and phosphorus are the primary nutrients that affect water quality in streams in the midwestern USA and high concentrations of these nutrients tend to increase algal biomass. However, how nutrients interact with physical controls in regulating algal biomass is not well known in agricultural streams. Eighteen streams in east-central Illinois (USA) were sampled during June and September 2003 to analyze factors possibly regulating algal biomass. Additionally, two shaded and two non-shaded sites in the Embarras River in east-central Illinois were sampled intensively from June to December 2003. Both sestonic and periphytic chlorophyll-a (chl-a) were analyzed, and periphytic chl-a was assessed on natural substrata and unglazed ceramic tiles. Although high concentrations of nutrients were found in these streams (mean total P = 0.09–0.122 mg l⁻¹ and mean NO₃-N=4.4–8.4 mg l⁻¹), concentrations of sestonic chl-a were low among all sites and both sampling periods (<18 mg m⁻³, median values of 5 and 3 in June and September, respectively). Filamentous algae were an important component of the algal communities in streams with stable substrata. Periphytic chl-a was generally not related to the concentration of N or P in the water column, and in non-shaded streams periphyton appeared at times to be light-limited due to turbid water. Turbidity was found to be an important factor controlling chl-a on ceramic tiles across the 18 sites and for the Embarras River sites; chl-a decreased exponentially in concentration (132–0 mg m⁻²) as turbidity increased from 4 to 39 NTU (r ² = 0.80). In general, the interaction between hydrology and light (turbidity) likely controlled algal biomass in these nutrient-rich, agricultural streams.     

Changes in Phytoplankton Biomass and Nutrient Quantities in Sea Ice as Responses to Light/Dark Manipulations during different Phases of the Baltic Winter 2003

The response of Baltic Sea ice communities to changing light climate was studied in three subsequent 3 week in situ experiments on the SW coast of Finland. The investigation covered three different winter periods, short day with low solar angles leading to limited light in the ice, late winter with deep snow cover and early spring with melting snow and increasing light availability. The experimental setup consisted of transparent (no snow) and completely darkened (heavy snow cover) plexiglass tubes in which the ice cores were incubated in situ from 1 to 2 weeks. Changes in the concentrations of inorganic nutrients (NO₃⁻-–N, PO₄³⁻-–P, SiO₄⁻-–Si) and chlorophyll-a concentration in the phytoplankton community composition were recorded as responses to different light manipulations. Changes in inner ice light intensity in untreated ice as well as the temperature both in air and ice were recorded over the entire study period. Increased irradiance in late winter/early spring and during meltdown affected the chlorophyll-a amount in the sea ice. During these periods the phytoplankton community in the top layers decreased possibly as a consequence of photo-acclimation. Closer to the bottom of the ice, however, the increased inner ice light intensity induced algal growth. Complete exclusion of light stopped the algal growth in the whole ice column. Darkening the ice cores also slowed down the ice melting opposite to accelerated melting caused by increased light. The significant differences found in nutrient concentrations between the light and dark treatments were mostly explicable by changes in algal biomass. No obvious changes were observed in the phytoplankton community composition due to light manipulation, diatoms and heterotrophic flagellates dominating throughout the study period.     

Different responses to changes in phosphorus,P, among lakes. A study of slopes in chl-a = f(P) graphs

The least squares estimator of a linear regression coefficient _L will give an overall expression for the change in with x. In fresh water ecology, however, subgroups, % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaamiuaSGaci% 4Aaaaa!37BE!\[P\operatorname{k}\], of a parent population may have slopes which differ from the overall slope, _L. By constructing frequency histograms for the set of angles: Arctang % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaci4uaSGaam% yAaiaadQgaaaa!38AE!\[\operatorname{S} ij\],% MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaci4uaSGaam% yAaiaadQgaaaa!38AE!\[\operatorname{S} ij\]= para sa y and x% MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaaiikaiaadM% faliaadMgakiabgkHiTiaadMfaliaadQgakiaacMcacaGGVaGaaiik% aiaadIhaliaadMgakiabgkHiTiaadIhaliaadQgakiaacMcaaaa!42F0!\[(Yi - Yj)/(xi - xj)\], i < j, % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaamiEaSGaam% yAaOGaeyiyIKRaamiEaSGaamOAaaaa!3BAB!\[xi \ne xj\], peaks in the distribution may be identified and related to ecological phenomenon. To identify peaks we fit Gaussian distributions to the frequency histograms. For a set consisting of 142 observations of chlorophyll-a and total phosphorus (nutrient) concentrations (TP) from 16 lakes we found four Gaussian peaks corresponding to four subgroups, % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaamiuaSGaci% 4Aaaaa!37BE!\[P\operatorname{k}\]k = 1,4. One group identified a response of chl-a to changes in TP which correspond approximately to the average slope found by least square regression (the slope was 0.49). The second group consisted of steeper response than the average (1.28). A third group showed that there is an enhanced proportion of cases where chl-a does not respond to TP (zero slope, all the three deep lakes > 10 m, included in the date set contributed to this group). The size of the last group, spanning a wide range of slopes, suggested that about 30% of the inter annual changes in chl-a is unrelated to TP. The results are compared to result obtained by simple least squares regression and to the Theil non-parametric slope estimator.