Relationship between Photosynthesis and Plasmalemma Transport

Hansen, U.-P., Kolbowski, J. and Dau, H. 1987. Relationshipbetween photosynthesis and plasmalemma transport.—J. exp.Bot. 38: 1965–1981. The yield of chlorophyll-fluorescence (F), of oxygen evolution(PAS, photo-acoustic signal) and the light-induced changes ofplasmalemma potential (V) were measured in the presence of red(633 nm) background light with blue or far-red (720 nm) sinusoidallymodulated actinic light. The kinetic study based on curve-fittingof the measured frequency responses led to the following: theresponse of chlorophyll-fluorescence (F) to blue actinic lightcomprises four time-constants located at 2·7,50,400 and4000 s. Membrane potential (V) displays five time-constants:2· 50, 160±330, —1, 4000 s. These time-constantswere assigned to the reactions involved as follows: from thecomparison of the effects of blue and far-red actinic lightthe time-constants of 2·7 s and 400 s were related tothe plastoquinone pool and to the state-transition controller.The time-constant of 4000 s was not investigated. The complextime-constants of V are known to be related to the controllerof cytoplasmic pH from previous studies. The time-constant of50 s was common to V, F, and PAS. From the sign of the 50 scomponent in F and in PAS (q_E-component), the following modelof the light action on membrane transport was developed: theuptake of protons into the inner space of the thylakoids causesan alkalinization of the cytoplasm which slows down the plasmalemmaH₊-pump via a substrate effect. Key words: Chlorophyll-fluorescence, frequency responses, kinetic analysis, plasmalemma potential, photo-acoustic effect, proton flux, spinach, state-transitions, thylakoid membrane, time-constants     

温室油桃光合作用与生理生态因子的关系

以日光温室栽培4年生油桃品种'艳光'为试材,采用LI-6400光合仪测定了成熟期的叶片净光合速率(Pn)以及气孔导度(Gs)、蒸腾速率(Tr)、光合有效辐射(PAR)、叶片温度(Tl)和CO2浓度(Ca)等生理生态因子的日变化,并运用数理统计方法分析了Pn与各因子的关系,为温室油桃合理栽培选择提供理论依据.结果表明:'艳光'油桃成熟期叶片的Pn日变化为"双峰"曲线,最高峰出现在12:00左右,次高峰出现在16:00左右,存在"午休"现象,非气孔因素是导致"午休"现象的主要原因;Pn与Gs、Tr和Ca均存在显著的二次曲线关系,与PAR有显著双曲线关系,与Tl有显著"S"型曲线关系;温室油桃光合作用的最适温度为25℃,光饱和点和光补偿点分别为860和26 μmol·m-2·s-1;二氧化碳饱和点和补偿点分别为1 340和54 μmol/mol.因此,在本实验条件下,温室油桃果实成熟期应当保证充足水分供应,并在正午前后适当采取降温措施,尽量减轻叶片气孔导度和蒸腾速率下降及光合'午休'现象发生程度,保证正午光合作用的正常进行,提高油桃的光能利用率.     

The Rate of Photorespiration during Photosynthesis and the Relationship of the Substrate of Light Respiration to the Products of Photosynthesis in Sunflower Leaves

Single attached leaves of sunflower (Helianthus annus L. “Mennonite”) were supplied ¹⁴CO₂ of constant specific radioactivity in gas mixtures containing various CO₂ and O₂ concentrations. The ¹⁴CO₂ and CO₂ fluxes were measured concurrently in an open system using an ionization chamber and infrared gas analyzer.     

Source-Sink Relationship: Control of Photosynthesis in the Flag Leaf by the Panicle in Wheat

If the sink organs regulate the metabolic activity of the sourceorgans through the transmission of a signal and that signalis chemical in nature, then it should be present among the substancescoming out of the panicle. When flag leaf respiration and photosynthesisof wheat (Tritcum aestivum L.) cv. HP 1459 were measured withan infrared gas analyser after feeding the flag leaves withthe diffusate coming out of the panicle, it was observed thatwhile respiration was unaffected, photosynthesis was sharplyinhibited; the inhibition decreased with increasing age of thepanicle. Among the known plant growth regulators none couldsatisfy the requirements of this role with the exception ofabscisic acid which was found to inhibit photosynthesis by morethan 50% even at 10^–7 M, when respiration was affectedby only 8%. Apparently, abscisic acid inhibited photosynthesisthrough its effects on stomatal movement. (Received September 2, 1986; Accepted May 1, 1987)     

小麦叶片光合作用与其渗透调节能力的关系

在缓慢干旱条件下,小麦叶片渗透调节能力在一定范围内随胁迫程度的加剧而增加,而在快速干旱下,渗透调节能力丧失。小麦叶片通过渗透调节使光合速率和气孔导度对水分胁迫的敏感性降低,叶片维持较高的电子传递能力、RuBP羧化酶活性和叶绿体光合能量转换系统活性,并推迟了小麦叶片光合速率受气孔因素限制向叶肉细胞光合活性限制转变的时间。     

The Relationship Between the Rates of Carbon Transport and of Photosynthesis in Tomato Leaves

The rate of carbon transport based on the carbon balance overa 6-h period from a mature tomato leaf was measured overa rangeof net photosynthetic rates from 0.1 to 4.9 mg C dm^–2h^–1 under light flux densities from 4 to 140 W m^–2.A proportional relationship was demonstrated between the rateof carbon transport and carbon fixation when the carbon fixationrate was higher than 2 mg C dm^–2 h^–1.Below a carbonfixation rate of 1 mg C dm^–2 h^–1, the rate of carbonexport was maintained at 1 mg C dm^–2 h^–1 at theexpense of the breakdown of starch. A highly significant correlationwas observed between sucrose concentration and the rate of carbontransport. The sucrose concentration in the leaf appears tobe the factor controlling carbon export.     

The Relationship Between Photosynthesis and Tuber Growth in Solanum tuberosum L.

Measurements of the rate of photosynthesis of plants of Solanumtuberosum L. var. King Edward were made, using ¹⁴CO₂, at weeklyintervals throughout their growth in a controlled environment.Leaf area and dry weight of sections of the plant were alsodetermined. The results are discussed in relation to existingtheories that photosynthesis can be limited by carbohydrateaccumulation in the leaves, and stimulated by the initiationof tubers.     

光合单位的周转率与叶片光合速率的关系

利用闪光技术,测得若干高等植物叶片的光合单位为 1450～2050 Chl/CO_2。在含O_2 2％及CO_2浓度和光通量密度都饱和的条件下,测得叶片的连续光合速率为25～70μmol CO_2 m~(-2) s~(-1)。从上述数据及叶片叶绿素含量计算,得叶片光合单位的周转率已达80～160次/s。这样的周转速率与Emerson当初用改变闪光间暗间隔而测到的光合单位周转时间相似(即<10 ms/次,或>100次/s),从而证实了光合单位的周转与光合速率直接有关。这样的速率可称之为叶片光合的理论速率。     

祁连圆柏光合作用日变化特征及其与生理生态因子的关系

以吐鲁沟国家森林公园内百年左右树龄的祁连圆柏为对象,采用Li-6400光合作用测定系统,在自然环境条件下测定了其生长季内各月份生理生态因子和净光合速率的日变化,探讨祁连圆柏生理生态特征以及与其生理生态因子间的关系.结果表明,在整个生长季内,祁连圆柏的净光合速率日变化为双峰型曲线,其在10:00和16:00达到峰值,于14:00达到最小值,光合作用存在明显的午休现象",而月份之间的净光合速率表现为8月>5月>6月>7月.祁连圆柏的气孔限制值与净光合速率有相似的日进程,呈现出M"型的双峰曲线,分别在10:00和16:00达到峰值,而胞间CO2浓度日变化趋势与气孔限制值相反,表现出W"型双峰曲线.水分利用效率与净光合速率有相同的日变化趋势,其在整个生长季内表现为5月>8月>7月>6月.在整个生长季内,影响祁连圆柏光合变化的主要决定生理生态因子是胞间CO2浓度和气温,限制生理生态因子是气孔导度和光合有效辐射强度.     

Light-induced Changes in the Ultrastructure of Pea Chloroplasts in Vivo: Relationship to Development and Photosynthesis

Light-induced structural changes of chloroplasts and their lamellae were studied in leaves of Pisum sativum L., cv. Blue Bantam, using electron microscopy. Upon illumination of 14-day-old plants with 2000 lux, the chloroplasts decreased in thickness by about 23% with an accompanying increase in electron scattering by the stroma. Concomitantly, the average thickness of granal lamellae (thylakoids) decreased from 195 ± 4 angstroms in the dark to 152 ± 4 angstroms in the light, and this change was half-saturated at only 50 lux. Lamellar flattening at 50 lux and its reversal in the dark both had half-times of a minute or less. The thickness of a partition (a pair of apposed lamellar membranes) was 140 ± 9 angstroms in both the light and the dark, indicating that the observed light-induced change was in the volume enclosed within the thylakoid. The effect of illumination could be inhibited by various uncouplers of photophosphorylation but not by 3-(3, 4-dichlorophenyl)-1, 1-dimethylurea, suggesting that it depended on ATP (or its precursor). In the presence of 0.5 micromolar nigericin, the thickness of the granal lamellae increased in the light to 213 ± 3 angstroms; this may reflect an uptake of K⁺ into an osmotically responding space within the thylakoids.     

羌活光合作用日变化及其与生理生态因子的关系

以大田引种栽培的羌活植株为材料,用Li-6400便携式光合作用测定系统原位测定自然条件下生长的羌活孕蕾期叶片的光合速率光响应、净光合速率及生理生态因子日变化,并通过相关分析、通径分析和逐步回归分析探讨净光合速率与生理生态因子的关系.结果表明:(1)羌活净光合速率(P_n)、 蒸腾速率(T_r)、气孔导度(G_s)日变化均呈双峰曲线,其净光合速率具有典型的"午休"现象,并主要由非气孔限制因素造成;(2)影响羌活叶片Pn日变化的主要决定生理因子是G_s,主要限制因子是胞间CO_2浓度(C_i);主要决定生态因子是空气相对湿度(RH),限制因子是气温(T_a); G_s是影响净光合速率最重要的生理生态因子.研究发现,羌活有较强的适应弱光环境的能力,属于阳性耐荫植物,宜选择海拔和荫蔽度均较高的环境进行引种驯化栽培,以利于其生长和存活.     

Photosynthesis in the Archean Era

The earliest reductant for photosynthesis may have been H₂. The carbon isotope composition measured in graphite from the 3.8-Ga Isua Supercrustal Belt in Greenland is attributed to H₂-driven photosynthesis, rather than to oxygenic photosynthesis as there would have been no evolutionary pressure for oxygenic photosynthesis in the presence of H₂. Anoxygenic photosynthesis may also be responsible for the filamentous mats found in the 3.4-Ga Buck Reef Chert in South Africa. Another early reductant was probably H₂S. Eventually the supply of H₂ in the atmosphere was likely to have been attenuated by the production of CH₄ by methanogens, and the supply of H₂S was likely to have been restricted to special environments near volcanos. Evaporites, possible stromatolites, and possible microfossils found in the 3.5-Ga Warrawoona Megasequence in Australia are attributed to sulfur-driven photosynthesis. Proteobacteria and protocyanobacteria are assumed to have evolved to use ferrous iron as reductant sometime around 3.0 Ga or earlier. This type of photosynthesis could have produced banded iron formations similar to those produced by oxygenic photosynthesis. Microfossils, stromatolites, and chemical biomarkers in Australia and South Africa show that cyanobacteria containing chlorophyll a and carrying out oxygenic photosynthesis appeared by 2.8 Ga, but the oxygen level in the atmosphere did not begin to increase until about 2.3 Ga.     

Relationship between Carbon and Nitrogen Metabolisms in Photosynthesis. The Role of Photooxidation Processes

¹⁴CO₂ uptake in leaves of wheat plants (Triticum aestivum L.) fertilized by urea or Ca(NO₃)₂ (25 mol m^-3) was investigated. The Warburg effect (inhibition of ¹⁴CO₂ uptake by oxygen) under 0.03 vol. % CO₂ concentration was observed only in non-fertilized plants. Under 0.03 vol. % CO₂, the Warburg antieffect (stimulation of ¹⁴CO₂ uptake by oxygen) was detected only in plants fertilized by Ca(NO₃)₂. Under saturating CO₂ concentration (0.30 vol. %), the Warburg antieffect was observed in all variants. Under limitation of ribulose-1,5-bisphosphate carboxylase/oxygenase activity (0.30 vol. % CO₂ + 1 vol. % O₂), the rate of synthesis of glycollate metabolism products decreased in control and urea-fertilized plants but was enhanced in nitrate-fed plants. Hence, there was an activation of glycollate formation via transketolase reaction in fertilized plants, and the products of nitrate reduction function were oxidants in nitrate-fertilized plants whereas the superoxide radical played this role in urea-fertilized plants.     

干旱胁迫下景天植物光合作用与超微弱发光的关系

该试验以德景天幼苗为试材,设计PEG、PEG+H_2O_2、PEG+苯甲酸钠、蒸馏水(CK)4个处理,分析20%PEG模拟干旱胁迫、以及活性氧调控干旱胁迫下超微弱发光(UWL)和光合作用参数的变化特征及其两者的关系,为揭示UWL的产生及其来源提供理论依据。结果表明:(1)在PEG模拟干旱胁迫过程中,CK和PEG处理的德景天叶片的净光合速率(Pn)、蒸腾速率(Tr)、胞间二氧化碳浓度(Ci)、气孔导度(Gs)和叶绿素a(Chla)、叶绿素b(Chlb)、叶绿素总量(Chl)以及UWL强度均随处理时间呈下降趋势,但PEG处理的以上所有指标的下降较CK更快、降幅更大。(2)H_2O_2和苯甲酸钠调控PEG干旱胁迫过程中,PEG+H_2O_2、PEG+苯甲酸钠处理的Pn、Tr、Gs、Ci、Chla、Chlb、Chl以及UWL强度的变化趋势与PEG处理基本一致,均随处理时间呈下降趋势,但PEG+H_2O_2处理的上述光合指标和UWL强度均低于PEG处理,PEG+苯甲酸钠处理的光合指标和UWL强度却均高于PEG处理。说明苯甲酸钠处理具有缓解干旱胁迫并提高UWL强度的作用。(3)相关分析显示,干旱胁迫及活性氧调控干旱胁迫下UWL强度均与各光合指标呈显著正相关关系。研究发现,在干旱胁迫和活性氧调控干旱胁迫下德景天光合作用参数较CK均明显下降,UWL强度也随之明显降低;德景天叶片UWL强度随着光合作用效率的下降而降低,说明植物UWL的产生与其光合作用的强弱显著相关,植物UWL强度能够反映植物受到的逆境胁迫的程度。     

Differences between Wheat Genotypes in Specific Activity of Ribulose-1,5-bisphosphate Carboxylase and the Relationship to Photosynthesis

The in vitro ribulose-1,5-bisphosphate (RuBP) carboxylase activity per unit of leaf nitrogen was found to be 30% greater in Triticum aestivum than in T. monococcum. This was due to a higher specific activity of the enzyme from T. aestivum, as the amount of RuBP carboxylase protein per unit of total leaf nitrogen did not differ between the genotypes. The occurrence of higher specific activity of RuBP carboxylase is shown to correlate with possession of the large subunit derived from the B genome of wheat.

Despite the greater RuBP carboxylase activity per unit of leaf nitrogen in T. aestivum, the initial slopes of curves relating rate of CO₂ assimilation to intercellular p(CO₂) are similar in T. aestivum and T. monococcum for the same nitrogen content per unit leaf area. The similarity of the initial slopes is the result of a greater resistance to CO₂ transfer between the intercellular spaces and the site of carboxylation in T. aestivum than in T. monococcum.

     

植物光诱导延迟荧光与光合作用的内在关联性研究

光合作用是地球上最重要的化学反应,植物内源性光诱导延迟荧光是光合作用原初过程中光系统Ⅱ作用中心P680处电荷分离效率的内在探针。本文从实验上证明了延迟荧光的产生与光合作用存在本质必然的联系。实验结果表明,延迟荧光激发谱与光合作用谱存在着明显的相似,延迟荧光强度随着激发光源光强的变化表现出类似的光抑制现象,以及在室温条件下延迟荧光强度与光合速率有着很好的相关性。为从植物光诱导延迟荧光技术来研究植物的光合作用提供了重要的实验证据。     

Relationship between Photosynthesis and Chlorophyll Content during Leaf Senescence of Rice Seedlings

Photosynthetic oxygen evolution, chlorophyll contents and chlorophylla /b ratios of 3rd to 6th leaves of rice seedlings were measuredto examine whether or not inactivation of photosynthesis duringsenescence is related to loss of chlorophyll. Photosyntheticactivity decreased more rapidly than chlorophyll content duringleaf senescence; as a result, the lower the leaf position, thelower was the rate of oxygen evolution determined on the basisof chlorophyll. Chlorophyll ab ratio also decreased with advancingsenescence. Electrophoretic analysis revealed that the declinein chlorophyll ab ratio is due to more rapid degradation ofthe reaction center complexes than light-harvesting chlorophyllab proteins of photosystem II and that the photosystem I reactioncenter disappears in parallel with the inactivation of photosynthesis.A simple method was developed to estimate the amounts of chlorophylla associated with the reaction center complexes of the two photosystemsfrom the total chlorophyll contents and chlorophyll ab ratiosof leaves. Rates of oxygen evolution, determined on the basisof chlorophyll a bound to the reaction center complexes, remainedconstant throughout the course of senescence. Thus, inactivationof photosynthesis is closely related with loss of the reactioncenter complexes during leaf senescence of rice seedlings. Theresults suggest that loss of photosynthesis is mainly causedby loss of a functional unit of photosynthesis or by a decreasein the number of whole chloroplasts. (Received April 22, 1987; Accepted August 13, 1987)     

无患子光合生理日变化及其与生理生态因子的关系

以8年生无患子植株为试验材料,采用便携式Li-6400光合测定仪,分别于花期和果期测定了其净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、胞间CO2浓度(Ci)、空气温度(Ta)、叶片光合有效辐射(PAR)、空气相对湿度(RH)和空气CO2浓度(Ca)等参数的日变化,以揭示无患子光合特征及其与主要环境因子的关系。结果显示:(1)无患子Pn和Tr在花期和果期日变化均呈"M"型双峰曲线,并存在"光合午休"现象,第1和第2峰值分别出现在10:00和14:00左右。(2)无患子花期Pn、Tr、Gs、水分利用效率(WUE)日均值高于果期,而果期的Ci日均值高于花期。(3)在8:00~14:00,无患子花期光能利用效率(LUE)大于果期,并在10:00左右出现峰值,而在14:00~18:00其果期LUE大于花期。(4)无患子叶片Pn与Tr、Gs、PAR具有极显著正相关关系,且与Gs关系最为密切,与Ci呈极显著负相关关系,与Ca则表现为显著负相关关系。(5)影响无患子叶片Pn变化的决定生理因子是Gs和Tr,主要限制生理因子是Ci;主要决定生态因子是RH,主要限制生态因子是Ta,且Gs是影响无患子Pn的最重要的生理生态因子。研究表明,无患子对光强有较强的适应能力,是喜光植物,这为其人工林定向培育,以及花期和果期的管理提供了一定理论依据。     

Photosynthesis in Wood

The capacity for photosynthesis, measured as light enhanced ¹⁴CO₂ uptake, was demonstrated in the wood of first and fourth year twigs of Betula pendula Roth, Populus tremuloides Michx., Syringa vulgaris L., and Tilia Americana L, but not in Gleditschia triacanthos L. In Betula and Syringa, but not in the others, photosynthesis occurred in the secondary xylem as well as in the primary xylem.     

Photosynthesis in the seeds of chloroembryophytes

Depending on the presence or absence of chlorophylls in the embryo, angiosperms are divided into chloroembryophytes and leucoembryophytes. Synthesis of chlorophylls (Chl) in the chloroembryos starts in the globular stage, rises as the embryo is formed, and stops in the late phase of seed maturation. The seeds also contain carotenoids that participate in photosynthesis and act as ABA precursors. The chloroembryos contain photochemically active chloroplasts that contain all the main photosynthetic complexes at a necessary stoichiometric ratio. Dark reactions of photosynthesis in developing seeds are notable for the fact that the main source of carbon therein is sucrose arriving from the maternal plant. Therefore, function of chloroplasts mainly aims at production of NADPH and ATP that are spent on conversion of sucrose into acetyl-CoA and, subsequently, to fatty acids. The CO₂ fixation system involving Rubisco and/or phosphoenolpyruvate carboxylase operates in the chloroembryos. In the course of photosynthesis, oxygen is released, which prevents hypoxia and maintains seed respiration. In late stages of ripening, the seeds enter the state of dormancy, which is associated with dehydration, disintegration of photosynthetic apparatus, Chl breakdown, and transformation of chloroplasts into plastids filled with reserve nutrient substances. At the same time, very often Chl are not destroyed completely and their residues are present in mature seeds of numerous plant species.