A cell-based model for the photoacclimation and CO2-acclimation of the photosynthetic apparatus

We have developed a mathematical model based on the underlying mechanisms concerning the responses of the photosynthetic apparatus of a microalga cell which grows under constant incident light intensity and ambient CO₂ concentration. Photosynthesis involves light and carbon-fixation reactions which are mutually dependent and affect each other, but existing models for photosynthesis don't account for both reactions at once. Our modeling approach allows us to derive distinct equations for the rates of oxygen production, NADPH production, carbon dioxide fixation, carbohydrate production, and rejected energy, which are generally different. The production rates of the photosynthesis products are hyperbolic functions of light and CO₂ concentration. The model predicts that in the absence of photoinhibition, CO₂-inhibition, photorespiration, and chlororespiration, a cell acclimated to high light and/or CO₂ concentration has higher photosynthetic capacity and lower photosynthetic efficiency than does a cell acclimated to low conditions. This results in crossing between the two curves which represent the oxygen production rates and carbon fixation rates in low and high conditions. Finally, in the absence of photoinhibition and CO₂-inhibition, the model predicts the carbohydrate production rate in terms of both light intensity and CO₂ concentration.     

冷锻炼对甜椒叶片光合作用及其低温光抑制的影响

以冷敏感植物甜椒 (CapsicumannuumL .)抗冷性不同的两个品种为试材 ,利用CIRAS 1光合测定系统和FMS2调制式荧光仪 ,在控温控光条件下分析比较了冷锻炼苗与未经锻炼苗的叶片光合特性、叶绿素荧光参数对温度的响应。结果表明 ,随着温度的降低 ,无论是否经过锻炼 ,低温主要通过抑制碳同化能力来影响光合作用 ,并使光能过剩 ,导致低温光抑制。提高环境CO2 浓度以增强暗反应对光能的利用 ,低温光抑制减轻。 5d的亚适温锻炼过程中甜椒叶片已发生一定程度的光抑制 ,但锻炼苗叶片能在低温下维持较高的光系统II光化学效率(ФPSII)、光化学猝灭系数 (qP)和光适应下光系统II最大光化学效率 (Fv′/Fm)值 ;冷锻炼提高了两品种低温下对光抑制的抗性 ,而且对抗冷品种的作用效果更明显     

Light Stress and Oxidative Cell Damage in Photoautotrophic Cell Suspension of Euphorbia characias L

A photoautotrophic cell-suspension culture of Euphorbia characias L. grown at 70 [mu]mol photons m-2 s-1 was very sensitive to light stress: the gross photosynthesis measured by using a mass spectrometric 16O2/18O2 isotope technique showed a fast decrease at a rather low light intensity of 100 [mu]mol photons m-2 s-1, far below the photosynthetic saturation level. The contribution of activated oxygen species on photosystem II photoinhibition was examined for a given light intensity. A protective effect on gross photosynthesis was observed with 1% oxygen. When light stress was applied to a methyl viologen-adapted cell suspension, photoinhibition was reduced. When 50 [mu]mol L-1 methyl viologen was added, photoinhibition was slightly enhanced. These responses suggested an involvement of superoxide radicals in the photoinhibition process of E. characias photoautotrophic cells. The long-term (16 h) effects of photoinhibition were then studied. Aldehyde (malondialdehyde and 4-hydroxyalcenals) production resulting from lipid peroxidation was stimulated in long-term stressed cells. When 50 [mu]mol L-1 methyl viologen were added, increased aldehyde production was measured. Under 1% oxygen, the aldehyde production was comparable to that of nonstressed cells. The relationship among lipid peroxidation, light intensity, and net photosynthesis suggests that aldehyde production may result from cell death provoked by a prolonged energy deficit due to the inhibition of photosynthesis.     

Photoinhibition and P700 in the Marine Diatom Amphora sp

The marine diatom Amphora sp. was grown at a light intensity of 7.0 × 10¹⁵ quanta centimeter⁻² second⁻¹. Light saturation of photosynthesis for these cells was between 6.0 and 7.0 × 10¹⁶ quanta centimeter⁻² second⁻¹. At light intensities greater than saturation, photosynthetic ¹⁴CO₂ fixation was depressed, while P700 unit size (chlorophyll a concentration/P700 activity) increased and number of P700 units per cell decreased. After a 1-hour exposure of Amphora sp. to a photoinhibitory light intensity of 2.45 × 10¹⁷ quanta centimeter⁻² second⁻¹, there was a 45 to 50% decrease in the rate of ¹⁴CO₂ fixation relative to the rate at the culture light intensity. There also was a 25% increase in P700 unit size and a 30% reduction in the number of P700 units per cell but no change in total chlorophyll a concentration. Following this period of photoinhibition, the cells were returned to a light regime similar to that in the original culture conditions. Within 1 hour, both number of P700 units per cell and P700 unit size returned to levels similar to those of cells which were kept at the culture light intensity. The rates of photosynthesis did not recover as rapidly, requiring 2 to 3 hours to return to the rate for the nonphotoinhibited cells. Our results indicate that a decrease in P700 activity (with a resultant increase in P700 unit size) may be partially responsible for the photoinhibition of algal photosynthetic carbon dioxide fixation.     

Photosynthetic carbon metabolism in isolated pea chloroplasts: metabolite levels and enzyme activities

We report here that enzyme activation precedes the rise in metabolite levels, which appear to limit photosynthetic CO2 fixation during induction in pea leaf chloroplasts. Therefore light activation may be required for the build-up of photosynthetic intermediates and hence for photosynthesis in isolated chloroplasts. Analysis of metabolite levels and the known kinetic properties of the chloroplast enzymes indicates that the reductive pentose phosphate cycle is subject to control which fluctuates between several points during induction and when CO2 fixation is maximal. The transketolase-aldolase-catalyzed reactions around sedoheptulose-biphosphatase appear to provide a simple and effective primary control for photosynthetic CO2 fixation. When substrate levels and enzyme active site concentrations are taken into account, there is insufficient glyceraldehyde 3-phosphate dehydrogenase, aldolase, and transketolase activity to support photosynthetic CO2 fixation at observed rates. These results suggest that there may be direct transfer of glyceraldehyde 3-phosphate among these enzymes in the pea chloroplast.     

光合作用对光和二氧化碳响应的观测方法探讨

用便携式光合仪LI-6400观测自然条件下生长的盆栽蚕豆叶片光合作用对光和二氧化碳的响应发现：（1）用未经过光合诱导的叶片观测光合作用对光的响应会得到即使在全太阳光强下光合作用仍然不饱和的假象;（2）利用某些经验方程计算的饱和光强远低于实际观测值;（3）在观测光合作用对CO2的响应过程中,每一次CO2浓度变化都应当伴随一次光合仪的匹配步骤,否则所得结果偏差很大;（4）在不饱和光下观测光合作用对CO2的响应,会导致对叶片光合能力的低估。     

Photoinhibition of photosynthesis in the cyanobacterium Microcystis aeruginosa

We have examined characteristics of the photoinhibition of photosynthesis which occur in the unicellular cyanobacterium Microcystis aeruginosa, following exposure to photon fluence rates in excess of those required for growth. Photoinhibition occurs following exposure of cells to a photon fluence rate of 1,000 μmol m^-2 s^-1, which is manifested as a decrease in either light-limited CO₂ fixation or light-saturated CO₂-dependent O₂ evolution. The extent and rapidity of this photoinhibition is greatly enhanced under CO₂-depleted conditions. Experiments in which cultures were sparged with different gases indicate that photoinhibition is not an obvious consequence of elevated O₂ tensions, unlike the photooxidative bleaching of photosynthetic pigments. Comparative studies on the photoinactivation of CO₂-dependent O₂ evolution and of the methyl viologen-dependent Mehler reaction, in whole cells, indicate that a primary site of light damage is within the photosynthetic electron-transport reactions and that carbon fixation is initially unaffected.     

High Photosynthetic Capacity in a Shade-Tolerant Crassulacean Acid Metabolism Plant (Implications for Sunfleck Use,Nonphotochemical Energy Dissipation,and Susceptibility to Photoinhibition)

Aechmea magdalenae Andre ex Baker, a constitutive Crassulacean acid metabolism (CAM) plant from the shaded Panamanian rain forest understory, has a maximum photosynthesis rate 2 to 3 times that of co-occurring C3 species and a limited potential for photosynthetic acclimation to high light. Chlorophyll fluorescence measurements indicated that (a) compared with co-occurring C3 species, photosynthetic electron transport in A. magdalenae responded more rapidly to light flecks of moderate intensity, attained a higher steady-state rate, and maintained a lower reduction state of plastoquinone during light flecks; (b) these characteristics were associated with phase III CO2 fixation of CAM; (c) when grown in full sun, A. magdalenae was chronically photoinhibited despite a remarkably high nonphotochemical quenching capacity, indicating a large potential for photoprotection; and (d) the degree of photoinhibition was inversely proportional to the length of phase III. Results from the light fleck studies suggest that understory A. magdalenae plants can make more efficient use of sun flecks for leaf carbon gain over most of the day than co-occurring C3 species. The association between the duration of phase III and the degree of photoinhibition for A. magdalenae in high light is discussed in relation to the limited photosynthetic plasticity in this species.     

May photoinhibition be a consequence, rather than a cause, of limited plant productivity?

Photoinhibition in leaves in response to high and/or excess light, consisting of a decrease in photosynthesis and/or photosynthetic efficiency, is frequently equated to photodamage and often invoked as being responsible for decreased plant growth and productivity. However, a review of the literature reveals that photoinhibited leaves characterized for foliar carbohydrate levels were invariably found to possess high levels of sugars and starch. We propose that photoinhibition should be placed in the context of whole-plant source–sink regulation of photosynthesis. Photoinhibition may represent downregulation of the photosynthetic apparatus in response to excess light when (1) more sugar is produced in leaves than can be utilized by the rest of the plant and/or (2) more light energy is harvested than can be utilized by the chloroplast for the fixation of carbon dioxide into sugars.     

Photosynthetic acclimation to low temperature by western red cedar seedlings

Imposition of low, but above freezing, temperatures resulted in a gradual increase in the cold hardiness of western red cedar seedlings. This was associated with a decrease in the maximum rates of photosynthetic CO₂ fixation and O₂ evolution, and changes in chlorophyll a fluorescence transients which indicated that photoinhibition had occurred. Maximum photosynthetic rates declined approximately 40% during cold hardening. The leaves changed colour from green to red-brown during the hardening process. The colour change was due to the synthesis of large amounts of the carotenoid rhodoxanthin. Lutein levels doubled, while chlorophyll declined slightly. Dehardening resulted in the rapid recovery of photosynthesis to control levels, the rapid disappearance of rhodoxanthin, and the return of lutein levels to control. It is suggested that rhodoxanthin accumulation at low temperature functions to decrease the light intensity reaching the photosynthetic apparatus. The combination of photoinhibition and rhodoxanthin synthesis probably serves to protect the photosynthetic capacity of the seedlings at low temperature.     

Effect of Nitrogen Supply on the Photosynthetic Performance of Leaves from Coffee Plants Exposed to Bright Light

Although Coffea arabica L. grows naturally in shaded habitats,it can be cultivated under high light intensity, but not withoutsevere photoinhibition mainly during the period of transferfrom the nursery into the field. The present work examines someof the changes in the photosynthetic performance induced byexposure to high light and the possibility of using enhancednitrogen levels to overcome photoinhibition. For that purpose,young plants of Coffea arabica L. (cv. Catuai) grown in a shadedgreenhouse were treated with 0, 1 and 2 mmol of nitrogen and4 weeks later exposed to full solar irradiation, outside. Visible damage due to exposure to full sunlight appeared within2 d in all plants, resulting in a reduced photosynthetic leafarea and drastic shedding of leaves in the unfertilized plants.These effects were considerably less in plants with the highestN dose. After 130 d of exposure, there was 100% mortality inplants receiving no extra nitrogen, compared with 30% in theplants treated with 2 mmol nitrogen. Photosynthesis rates, leafconductance and transpiration presented minimum values after4 d of light stress. Large changes in the photosynthetic capacity(measured at high CO₂ concentration and high light intensity),quantum efficiency and fluorescence yield (F_v/F_m) indicate thatnet photosynthesis rate in the air had been reduced by bothstomatal closure and by changes at the photochemical level.All indicators show that N-fertilized plants were less affectedby photoinhibition. Key words: Coffee plant, nitrogen, photoinhibition, photosynthesis     

高浓度CO2引起的海水酸化对小珊瑚藻光合作用和钙化作用的影响

为了探讨CO2海底封存潜在的渗漏危险对于海洋生物的可能影响,以大型钙化藻类小珊瑚藻(Corallina pilulifera)为研究对象,在室内控光控温条件下,通过向培养海水充入CO2气体得到3种不同酸化程度的培养条件(pH 8.1、6.8和5.5),24h后比较藻体光合作用和钙化作用情况。结果显示:相对于自然海水培养条件(pH 8.1),在pH 6.8条件下培养的小珊瑚藻光合固碳速率得到了增强,而在pH 5.5条件下光合固碳速率则降低;随着酸化程度的增强,藻体的钙化固碳速率越来越低,在pH 5.5条件下甚至表现为负值[(-2.53±0.57)mg C g-1干重h-1];藻体颗粒无机碳(PIC)和颗粒有机碳(POC)含量的比值随着酸化程度的加强而降低,这反映了酸化对光合和钙化作用的综合效应。快速光反应曲线的测定结果显示:随着酸化程度的增强,强光引起的光抑制程度越来越强;在酸化条件下,藻体的光饱和点显著降低,但pH 6.8和5.5之间没有显著差异;低光下的电子传递速率在pH 8.1和6.8之间没有显著差异,pH 5.5培养条件下显著降低;最大电子传递速率在pH 6.8时最大,在pH 5.5时最低。以上结果说明,高浓度CO2引起的海水酸化显著地影响着小珊瑚藻的光合和钙化过程,不同的酸化程度下,藻体的光合、钙化反应不同,在较强的酸化程度下(pH 5.5),藻体的光合和钙化过程都将受到强烈的抑制,这些结果为认识CO2海底封存渗漏危险对海洋钙化藻类的可能影响提供了理论参考。     

Effects of Photoinhibition and Its Recovery on Photosynthetic Functions of Winter Wheat Under Salt Stress

Effects of photoinhibition and its recovery on photosynthetic functions of winter wheat ( Triticum aestivum L.) under salt stress were studied. The results showed that several parameters associated with PSⅡ functions, e.g. Fv/Fo 、 Fv/Fm and qP were not influenced by lower salt concentration (200 mmol/L NaCl) while CO2 assimilation rate decreased significantly. When exposed to higher salt concentration (400 mmol/L NaCl), PSⅡ functions were significantly inhibited which led to the decrease of carbon assimilation. These results suggest that different concentrations of salt stress affected photosynthesis by different modes. Salt stress made photosynthesis more sensitive to strong light and led to more serious photoinhibition. Under lower concentration of salt stress, the QB-non-reductive PSⅡ reaction centers formed at the beginning of photoinhibition could be effectively used to compose active PSⅡ reaction center (RC) and repair the reversible inactivated PSⅡ RC. Under higher concentration of salt stress, PSⅡ reaction centers were seriously damaged during photoinhibition, the QB-non-reductive PSⅡ RC could only be partly effective at the early time of photoinhibition, thus led to the accumulation of QB-non-reductive PSⅡ RC in the course of restoration under dim light.     

转C4光合酶基因水稻的CO2交换和荧光特性

用转PEPC、PPDK、NADP-ME、PEPC+PPDK酶基因水稻(Oryza sativa L.)及原种为材料 ,研究了光合作用对光照、温度、CO2的响应和光抑制条件下的叶绿素荧光特性,结果如下: 1.转C4光合酶基因水稻的饱和光合速率比原种高,其中转PEPC、PEPC+PPDK双基因水稻的光饱和点比原种高200 μmol*m-2*s-1,饱和光合速率比原种分别高51.6%和 58.5%;转PEPC基因水稻的羧化效率比原种高49.3%,CO2补偿点降低26.2%;在高温(35 ℃)下,转PEPC基因水稻的光合速率比原种高17.5%.2.经光抑制处理8 d后,转PEPC、PEPC +PPDK酶基因水稻的PSⅡ光化学效率(Fv/Fm)和光化学猝灭(qP)下降20%- 30%,非光化学猝灭(qN)增加了约30%;但原种的Fv/Fm和qP下降了5 0%多,qN变化不明显,表明转C4光合基因水稻耐光抑制能力增强.这些结果为用生物技术提高水稻光合效率研究提供了新的依据和途径.     

Light-dependent induction of strongly increased microalgal growth by methanol

Low methanol concentrations (about 0.5% v/v) induce biomass production in cultures of the unicellular green alga Scenedesmus obliquus by more than 300%, compared to controls without this solvent. This effect on the microalgal growth was found to be dependent on the solvent concentration, the packed cell volume (PCV), light intensity and light quality. It could be shown that methanol addition leads to a decrease in size of the light harvesting complex (LHC) on the basis of chlorophylls and proteins, and thus to changes in structure and functioning of the photosynthetic apparatus. These alterations lead to enhanced photosynthesis and respiration rates. The action of methanol on the photosynthetic apparatus is thus comparable to the effect of enhanced CO(2) concentrations. These findings support the previously proposed pathway for methanol metabolization with CO(2) as the final product. We conclude that the subsequent assimilation of the increased CO(2) amounts by the Calvin-Benson cycle is a possible explanation for the methanol-mediated increase in biomass production in terms of PCV. The methanol effect is observed only in the light and in the presence of a functioning photosynthetic apparatus. Preliminary action spectra suggest that the primary photoreceptor is a chlorophyll-protein complex with two absorption maxima at 680 and 430 nm, which may possibly be attributed to the reaction center of photosystem II (PSII).     

Photoacclimation and the effect of the symbiotic environment on the photosynthetic response of symbiotic dinoflagellates in the tropical marine hydroid Myrionema amboinense

Symbiotic dinoflagellates of the genus Symbiodinium and residing in the tropical hydroid Myrionema amboinense acclimate to low photon flux associated with low light 'shade' environments by increasing the amount of photosynthetic pigments per algal cell. The photosynthetic light intensity (PI) curves suggested that the low-light pigment response involved an increase in the number of photosynthetic units (PSU) in the chloroplast in addition to any increases in PSU size. Comparisons of light-dependent portion of the P-I curves of freshly isolated zooxanthellae (FIZ) with those from symbionts within the intact animal suggest that the host cell environment reduced average light levels reaching the symbiotic algae by more than half. This phenomenon may protect the algae from photobleaching of pigments and/or photoinhibition of photosynthesis at high light intensities present in shallow water habitats. In addition, maximum photosynthesis (P(max)) of symbionts removed from the host cell was higher than that recorded from dinoflagellates in the intact association, suggesting that the availability of carbon dioxide for photosynthesis may be limited in the intact hydroid. Shaded polyps contained fewer zooxanthellae and had less tissue biomass (measured as protein) than unshaded polyps. However symbionts from shaded polyps acclimated to the low light intensities by increasing chlorophyll levels and photosynthetic rates. The higher photosynthetic rates may have resulted from increased availability of carbon dioxide associated with lower symbiont density. Calculations of the contribution of zooxanthellae carbon to the host animal's respiratory demand (CZAR) showed that zooxanthellae from shaded polyps living in the field potentially provide about the same amount of carbon to their host as zooxanthellae from polyps living in the field in unshaded high light intensities.     

Light adaptation and the role of autotrophic epiphytes in primary production of the temperate seagrass, Zostera marina L.

Photosynthetic features of Zostera marina L. and its autotrophic epiphyte community were investigated in a population inhabiting a shallow (1.3 m depth) water meadow in Great Harbor, Woods Hole, MA (U.S.A.). Photosynthesis versus irradiance (P-I) relationships were measured with respect to leaf age determined by the leaf position in the shoot bundle and by location of the tissue along the leaf axis. Therefore both age and light intensity gradients along the leaf axis were considered. The maximum photosynthesis (P_max) per dm² typically increased nearly two-fold along the leaf axis from leaf bases to apices. Photosynthetic rate on a chlorophyll (Chl) basis did not increase as dramatically along the leaf axis, and rates were usually lowest in tissues with the highest Chl content. The P-I relationships of leaves of different ages did not reveal photoinhibition even at light intensities > 1400 μE • m⁻² • s ⁻¹. Furthermore, no photoinhibition was observed in tissues from leaf blade bases, which never experienced high light levels (> 500 μE • m ⁻² • s⁻¹) in situ in Great Harbor. The initial slopes of the P-I curves and light compensation and saturation values varied along the leaf axis in relation to in situ light intensity gradients and in relation to leaf or tissue age. It appeared that leaf and/or tissue age was more important than light environment in determining P-I responses. The contribution of the autotrophic epiphyte community on Z. marina leaves to total photosynthesis per dm² was between 27 and 50%, and between 10 and 44% per mg chlorophyll. These levels of epiphyte photosynthesis can double the primary production of Z. marina leaves. No detrimental effects of epiphyte cover were realized in leaf maximal photosynthesis or P-I relationships. Non-epiphytized leaves and leaves from which epiphytes were removed showed essentially identical photosynthetic features. Light intensity and age gradients along the leaf axis control both the photosynthetic performance of the leaves and epiphyte biomass and photosynthesis.     

Study of Diurnal Changes in Photosynthetic Rate and Quantum Efficiency of Grapevine Leaves and Their Utilization in Canopy Management

Photosynthetic rate and quatum efficiency of grapevine (Vitis vinifera L. cv. Sauvignon blanc) leaves were measured under the field with ample soil water supply, and in phytotron with ample supply of water and mineral nutrients, constant air humidity and CO2 concentration, and optimum air temperature, respectively. Under field conditions CO2 assimilation quantum efficiency of leaves reached its maximum in the morning, which was followed by continuous decrease and midday depression. The leaves intercepting more light energy in the morning showed a higher quantum efficiency. Those leaves subjected continuously to strong irradiance exhibited a more obvious and longer midday depression. Reduction of leaf light interception around midday could reduce midday depression. Shaded leaves had a higher quantum efficiency than leaves under direct sunlight. The diurnal changes in photosynthetic rate and quantum efficiency of leaves were shown to be closely related to the variations in mesophyll resistance to CO2. In phytotron experiments the photosynthetic quantum efficiency of leaves was reduced after a certain period of illumination not only at 1200 μmol · m-2 · s-1 PFD, higher than the saturating light of vine leaves (≈1000 μmol · m-2 · s-1), which was caused by "photoinhibition”, but also at 800 and 200μmol · m-2 · s-1, which was similar to "photoinhibition”. But photosynthetic quantum efficiency of leaves exposed continuously to a very weak PFD (100 μmol · m -2 · s-1) remained contant. The diurnal changes in mesophyll resistance to CO2 of vine leaves could be partly related to photoinhibition. It is considered that, under field conditions without soil water limitation, midday depression of vine leaf photosynthesis could be a result of an increase of the mesophyll resistance induced by multiple effects of strong light, high temperature and low humidity. A higher light interception by canopy plane in the morning may be advantageous to exploit higher photosynthetic potentiality of leaves, but a lower light interception in the middle of day may reduce midday depression. The north-south orientation plane can provide optimum light regime and improve photosynthetic environment in vineyards.     

柑橘属光合作用的环境调节

光合机构的运转受环境影响很大,与柑橘的生长发育、产量和品质密切相关.结合我们的工作,综合论述了柑橘光合作用环境调节的研究进展.强光和紫外光导致光合作用下降与PSⅡ反应中心失活有关,光呼吸和叶黄素循环对光合机构有保护作用.温度胁迫下,光合作用下降主要是RuBPCase活性下降和PSⅡ反应中心失活引起,品种间存在差异.轻度水分胁迫引起的光合作用下降是气孔限制的结果,而严重水分胁迫导致光合作用的非气孔限制.提高CO₂浓度,能够促进柑橘的光合作用,进而促进柑橘的生长和提高其品质.阐述了N、P、S、Fe等矿质元素调节光合作用的机理及盐胁迫对光合作用的影响,指出了今后柑橘光合作用的研究方向.