基于FvCB模型的几种草本和木本植物光合生理生化特性

为研究不同生活型植物的光合能力及其叶片光合机构,采用直角双曲线修正模型和C3植物FvCB模型对7种木本植物和4种草本植物的CO₂响应曲线进行拟合,并对不同木本植物、不同草本植物和2种生活型植物的最大净光合速率(P_{n max})、Rubisco酶最大羧化速率(V_{c max})、最大电子传递速率(J_max)、光合暗呼吸速率(R_d)和叶肉阻力(r_m)等参数进行比较.结果表明: 7种木本植物P_{n max}大小顺序为乌桕、苎麻>润楠、海桐>青冈、苦槠、娜塔栎;乌桕、苎麻、润楠和海桐的V_{c max}显著大于青冈和苦槠;J_max大小顺序为乌桕>苎麻、海桐>娜塔栎、苦槠和青冈;润楠和苦槠的r_m显著大于乌桕、海桐和苎麻.商陆的P_{n max}显著大于藿香蓟和土牛膝;4种草本植物的V_{c max}无显著差异;商陆的J_max显著大于藿香蓟;龙葵和土牛膝的r_m显著大于藿香蓟;商陆的R_d显著大于藿香蓟和土牛膝.木本植物的P_{n max}、V_{c max}、J_max和r_m光合参数均显著大于草本植物,但二者的R_d无显著差异.不同物种之间以及2种生活型植物光合能力的差异主要是由叶片内部Rubisco酶羧化能力、电子传递能力和叶肉阻力等差异引起的.     

基于结构方程模型分析森林草原带草本物种丰富度对景观因子的响应

受损景观是由不同比例的植被斑块组成的镶嵌体,阐明植被斑块的景观结构特征对物种多样性的影响,有助于提高受损景观物种多样性保护。在塞罕坝自然保护区选取38个天然植被斑块,其中包括12个草地、11个灌木林和15个天然次生林。根据光合作用的不同途径将草本物种划分为C_3和C_4功能群。选取斑块面积、形状指数、隔离度指数及每个斑块500 m缓冲区内的森林和草地比例作为景观因子。通过结构方程模型探讨C_3、C_4草本物种丰富度与景观因子的相互关系。斑块面积(2.18—74.06 hm~2)与C_3、C_4草本及总物种丰富度分布格局均具有显著的正相关关系(P0.05),且对C_3草本的影响最大;形状指数(1.06—3.11)、隔离度指数(33.51—327.65)对C_3、C_4草本及总物种丰富度影响不显著(P0.05);C_3、C_4草本及总物种丰富度与毗邻斑块草地比例(4.20%—64.95%)呈正相关,而与森林比例(35.05%—95.80%)呈负相关。研究区斑块面积和毗邻斑块植被构成是影响C_3、C_4草本植物的主要因素。在破碎化景观中保存面积大的天然植被斑块及提高毗邻斑块草地比例是保护关键C_3、C_4草本植物物种的有效途径。     

干旱胁迫对2种光合类型C4荒漠植物叶片光合特征酶和抗氧化酶活性的影响

以荒漠C₄草本植物蔷薇猪毛菜(NADP苹果酸酶型,NADP-ME）和粗枝猪毛菜（NAD苹果酸酶型,NAD-ME）为研究对象,采用盆栽控水试验设置正常供水和轻度、中度、重度干旱处理（土壤含水量分别为田间持水量80%、60%、45%和35%）,通过测定不同程度干旱胁迫下叶片含水量、C₄光合特征酶和抗氧化酶活性等指标,探讨不同类型C₄荒漠植物光合特征酶和抗氧化系统对干旱逆境的适应机制。结果显示：（1）2种植物叶片含水量均随干旱胁迫的加剧不同程度降低。（2）叶片磷酸烯醇式丙酮酸羧化酶（PEPC）活性在中度干旱胁迫下显著增加而在重度干旱胁迫下急剧下降;蔷薇猪毛菜NAD-ME活性和粗枝猪毛菜NADP-ME活性都很低,且它们基本不受干旱胁迫的影响;随干旱胁迫的加剧,蔷薇猪毛菜NADP-ME活性呈下降趋势,而粗枝猪毛菜NAD-ME活性先显著增加而在重度干旱胁迫下显著降低。（3）随着干旱胁迫的加剧,叶片超氧化物歧化酶（SOD）活性呈下降趋势,过氧化物酶（POD）活性在不同程度干旱胁迫下均有不同程度增加;过氧化氢酶（CAT）活性在中度干旱胁迫下均有不同程度的增加,但在重度干旱胁迫下蔷薇猪毛菜CAT活性降低,而粗枝猪毛菜CAT活性显著增加;丙二醛（MDA）含量随干旱胁迫的加剧均有不同程度的增加。研究认为,一定程度干旱胁迫下,2种荒漠植物的PEPC活性均有增加;不同光合类型C₄植物叶片脱羧酶（NADP-ME和NAD-ME）对干旱胁迫的响应有明显的差异。POD和CAT是这两种C₄植物适应干旱胁迫的主要抗氧化酶,但蔷薇猪毛菜CAT在重度干旱胁迫下没有起到积极保护作用。     

改进指数模型对紫茉莉光合-光响应及CO2响应适用性研究

五种模型分别运用于紫茉莉的光合—光响应及CO₂响应曲线的拟合,研究其光合效率参数的变化,探讨紫茉莉光合—光响应及CO₂响应的最适模型。结果表明:(1)紫茉莉的光合—光响应及CO₂响应改进指数模型拟合R²均为0.999,拟合效果优于非直角双曲线、直角双曲线和直角双曲线修正模型。其饱和光强和最大净光合速率分别为797.299和7.879 μmolCO₂·m^-2·s^-1,饱和CO₂浓度和最大光合能力分别为1 264.447和16.783 μmol CO₂·m^-2·s^-1,均与实测值最接近;(2)五个模型拟合和预测的均方误差(MSE)、平均绝对误差(MAE),都是改进指数模型小于其他模型。改进指数模型为紫茉莉光合—光响应及CO₂响应曲线的最佳模型,实验结果可为紫茉莉的生理生态应用研究提供参考。     

基于FvCB模型分析盐分胁迫对棉花叶片光合作用的影响

为深入理解叶片光合特性对盐胁迫的响应机理,以棉花为试验材料,设置5个盐分(NaCl)浓度处理:0(CK)、50、100、150和200 mmol·L^-1,利用FvCB模型分析盐胁迫对棉花幼苗叶片光合特性的影响。结果表明:与CK相比,50和100 mmol·L^-1盐分处理增加了棉花叶片的最大羧化速率(V_{c max})和最大电子传递速率(J_max),但150和200 mmol·L^-1盐分处理显著降低了V_{c max}和J_max。叶片净光合速率(P_n)、叶肉导度(g_m)和暗呼吸速率(R_d)随盐分浓度升高而下降;与CK相比,50和100 mmol·L^-1盐分处理对g_m无显著影响,但P_n和R_d显著降低。150和200 mmol·L^-1盐分处理明显降低了P_n、g_m和R_d,且与0、50和100 mmol·L^-1盐分处理间存在显著差异;利用FvCB模型模拟了不同盐分胁迫下叶片净光合速率。与不考虑g_m的模拟结果相比,考虑g_m提高模拟值和实测值间的决定系数,并降低了平均绝对误差。棉花幼苗耐盐阈值为100～150 mmol·L^-1,随盐分浓度的增加,光合限制因素由叶肉因素转变为光合机构受损;引入g_m可以提高FvCB模型的模拟精度。     

美国黑核桃实生苗单叶片光合作用生理生态模型的建立与验证

在对叶片光合过程机理分析的基础上,结合数学分析方法,建立了模拟美国黑核桃单叶片光合作用的机理模型,该模型包含了叶片光合作用的限速的生化过程和气孔调节因素,对光合作用一气孔导度的耦合模型进行了简化,使之既便于应用,又能较准确地反映田间条件下的情况,并就光合作用对环境因子(太阳辐射、温度及CO2浓度等）变化的响应特征及响应的合理性进行了分析,使用整个生长季实测的叶片生理数据及生态环境数据对所建模型进行了验证,结果表明,模型可以较准确地模拟田间美国黑核桃叶片的光合速率;确定了美国东部黑核桃、北加州黑核桃等两种黑核桃单叶片光合作用与环境因子互作的数量美系。     

不同温度及二氧化碳浓度下培养的龙须菜光合生理特性对阳光紫外辐射的响应

为了研究不同温度及CO₂浓度下培养的大型海藻对紫外辐射的生理学响应,选取龙须菜(Gracilaria lemaneiformis)作为实验材料。实验设置两个温度梯度(20 ℃和24 ℃),两种CO₂浓度(390 μL/L和1000 μL/L)以及3种辐射处理,即可见光(PAR)处理(滤除紫外线A(UV-A)、紫外线B(UV-B),400-700 nm)、可见光加紫外线A(PA)处理(滤除UV-B,320-700 nm)、PAB处理(全波长辐射280-700 nm)。结果表明,酸化、升温以及紫外辐射处理都未影响大型经济红藻龙须菜的叶绿素a和类胡萝卜素的含量。然而紫外辐射处理显著降低了龙须菜的有效光化学效率,其抑制水平在酸化处理的藻体中更为显著,并且随着温度的上升而进一步加剧;酸化与温度耦合使藻体对紫外辐射的敏感性增加,导致其较低的修复速率以及较高的损伤速率。     

曼地亚红豆杉对甲醛胁迫的生理响应

以一年生曼地亚红豆杉(Taxus media cv.hicksii)扦插苗为材料,采用密闭箱静态熏气法,研究不同甲醛（CH₂O）浓度(0、5、10、20和40 mg·m^-3)和熏气时间（1、3、5、7 d）对曼地亚红豆杉的生理响应。结果显示:（1）在5~20 mg·m^-3CH₂O浓度下,曼地亚红豆杉叶片均无受害症状,在40 mg·m^-3CH₂O熏气1 d时,叶片开始出现受害症状,并随时间的延长逐渐加重;（2）随着CH₂O浓度的增加和熏气时间的延长,叶片MDA、Pro含量和相对电导率皆呈增加趋势,SS含量表现为先升后降,但仍显著高于对照;（3）在5 mg·m^-3CH₂O处理下,叶片SOD、CAT、PPO和GR作为第一道防线共同作用以清除过多的活性氧,其中PPO最为敏感;在10、20 mg·m^-3CH₂O处理下,SOD、POD、CAT、PPO、APX和GR共同作用加快对活性氧的清理;在40 mg·m^-3CH₂O浓度下,各酶的活性均受到抑制,其中APX、PPO和GR活性显著低于对照,而SOD、POD和CAT活性仍显著高于对照。研究表明,在中低CH₂O浓度（5~20 mg·m^-3）处理下,曼地亚红豆杉主要通过合成渗透调节物质和活性氧自由基的酶促清除机制共同作用来适应逆境,在40 mg·m^-3CH₂O浓度下,APX、PPO、GR活性受到显著抑制,细胞膜过氧化程度加剧,植物叶片受到伤害;在CH₂O浓度低于20 mg·m^-3时,曼地亚红豆杉通过自身的应激保护系统来维持正常的生理活动,表现出较强的CH₂O耐受性。     

C4荒漠植物猪毛菜与木本猪毛菜的叶片解剖结构及光合生理特征

 为了比较C₄荒漠植物猪毛菜(Salsola collina)和木本猪毛菜(S. arbuscula)的抗旱结构和适应环境的光合作用特征, 在二者混生的群落中, 选择代表性植株, 采集叶片进行叶片解剖结构分析, 在自然条件下测定了二者叶片的气体交换参数。研究结果表明：猪毛菜叶片具表皮毛, 具有更发达的薄壁贮水组织;木本猪毛菜叶片具有更厚的角质层, 表皮下有1层下皮细胞, 其栅栏组织细胞较长, 排列更紧密。猪毛菜的净光合速率明显高于木本猪毛菜, 日平均值分别为21.5和15.7 μmol CO₂·m^–2·s^–1。猪毛菜的蒸腾速率也明显高于木本猪毛菜, 日平均值分别为14.9和10.2 mmol·m^–2·s^–1。猪毛菜和木本猪毛菜的水分利用效率的日平均值分别为1.39和1.53 μmol CO₂·mmol^–1 H₂O, 特别是在14:00时分别为1.61和2.30 μmol CO₂·mmol^–1 H₂O, 木本猪毛菜高出猪毛菜约42%。猪毛菜的光补偿点低于木本猪毛菜, 而光饱和点和光量子效率较高, 具有更低的CO₂补偿点。这表明：二者的旱生结构不同, 木本猪毛菜具有更显著的荒漠植物特征;在适于二者混生的环境下, 猪毛菜比木本猪毛菜的光合能力更强, 而木本猪毛菜的水分利用效率更高。     

半干旱雨养区小麦叶片光合生理生态特征及其对环境的响应

分析了黄土高原半干旱雨养农业区田间春小麦叶片光合生理生态特征及其对环境因子的响应。结果表明,天气晴朗时,净光合速率日变化呈典型的双峰曲线,有“午休”现象,上午明显高于下午,且不同生育期峰值出现的迟早不同。蒸腾速率日变化呈不明显的双峰型,其出现最大值的时间晚于净光合速率出现最大值的时间。在生长季节,叶片净光合速率、蒸腾速率和气孔导度均受到多个环境因子的共同影响。不同时期,起主导作用的环境因子不同,且同一个因子对几个生理指标的影响程度和强度都有差异,其中,光合有效辐射是对蒸腾速率影响最强烈的环境因子,湿度对光合作用的影响大于温度。受环境因子制约最为显著的生理指标是叶片的蒸腾速率和气孔导度。     

Carbon isotope composition as a tool to control the quality of herbs and medicinal plants

Isotope screening is a simple test for determining the photosynthetic pathway used by plants. The scope of this work was to classify the photosynthetic type of some herbs and medicinal plants through studies of the carbon isotope composition (δ¹³C). Also, we propose the use of carbon isotope composition as a tool to control the quality of herbs and medicinal plants. For studies of δ¹³C, δ¹³C‰ = [R (sample)/R (standard) − 1] × 10⁻³, dry leaves powdered in cryogenic mill were analyzed in a mass spectrometer coupled with an elemental analyzer for determining the ratio R = ¹³CO₂/¹²CO₂. In investigation of δ¹³C of 55 species, 23 botanical families, and 44 species possessed a C₃ photosynthetic type. Six species found among the botanical families Euphorbiaceae and Poaceae were C₄ plants, and 5 species found among the botanical families Agavaceae, Euphorbiaceae, and Liliaceae possessed CAM-type photosynthesis. Carbon isotope composition of plants can be used as quality control of herbs and medicinal plants, allowing the identification of frauds or contaminations. Also, the information about the photosynthetic type found for these plants can help in introducing and cultivating exotic and wild herbs and medicinal plants.     

Effects of polyploidy on photosynthesis

In polyploid plants the photosynthetic rate per cell is correlated with the amount of DNA per cell. The photosynthetic rate per unit leaf area is the product of the rate per cell times the number of photosynthetic cells per unit area. Therefore, the photosynthetic rate per unit leaf area will increase if there is a less than proportional increase in cell volume at higher ploidal levels, or if cell packing is altered to allow more cells per unit leaf area. In autopolyploids (Medicago sativa, C₃ species, and Pennisetum americanum, C₄ species) there is a doubling of photosynthesis per cell and of cell volume in the tetraploid compared to the diploid. However, there is a proportional decrease in number of cells per unit leaf area with this increase in ploidy such that the rate of photosynthesis per leaf area does not change. There is more diversity in the relationship between ploidal level (gene dosage) and photosynthetic rates per unit leaf area in allopolyploids. This is likely to reflect the effects of natural selection on leaf anatomy, and novel genetic interactions from contributed genomes which can occur with allopolyploidy. In allopolyploid wheat (C₃ species) a higher cell volume per unit DNA at the higher ploidal level is negatively correlated with photosynthesis rate per unit leaf area. Although photosynthesis per cell increases with ploidy, photosynthesis per leaf area decreases, being lowest in the allohexaploid, cultivated bread wheat (Triticum aestivum). Alternatively, doubling of photosynthetic rate per cell with doubling of DNA, with apparent natural selection for decreased cell volume per unit DNA, results in higher rates of photosynthesis per leaf area in octaploid compared to tetraploid Panicum virgatum (C₄) which may be a case of allopolyploidy. Similar responses probably occur in Festuca arundinacea. Therefore, in some systems anatomical factors affecting photosynthesis are also affected by ploidal level. It is important to evaluate that component as well as determining the effect on biochemical processes. Current information on polyploidy and photosynthesis in several species is discussed with respect to anatomy, biochemistry and bases for expressing photosynthetic rates.Abbreviations Chl chlorophyll - RuBPC ribulose-1,5-bisphosphate carboxylase     

毛竹光合作用对环境因子的季节响应

采用Li-6400测定毛竹光合作用对光照强度、CO2浓度、温度和湿度等环境因子响应的季节变化,结果表明:毛竹最大净光合速率、光补偿点、光饱和点、光合量子效率的年均值分别为7.30、19.15、1075mmol.m-2.s-1,0.032;最大净光合速率夏季>秋季>冬季>春季;春季的光补偿点最高,夏季次之,而秋季和冬季均较小;光饱和点与光合量子效率的季节变化均为秋季>夏季>冬季>春季。毛竹CO2补偿点、CO2饱和点、羧化效率的年均值分别为73.52、1500μmol.mol-1,0.033。CO2补偿点春季>冬季>秋季>夏季;CO2饱和点春季>秋季>夏季>冬季;羧化效率夏季>秋季>冬季>春季。毛竹光合最适温度均在20～30℃,光合最适温度在春、秋季与实验前3天最高气温的平均值十分接近,而夏、冬季与测定前10天的最高气温平均值较为接近,光合最适温度在春、秋两季相当,夏季稍高,冬季最低。光合最适湿度为40%～65%,季节变化趋势:秋季>夏季>冬季>春季。总体而言,毛竹光合作用对环境因子的季节响应与环境因子的季节变化、叶片的生理活性密切相关。     

Photosynthetic characteristics of mesophyll eells isolated from sunflower (helianthus annuus L.) leaves

Mesophyll cells were isolated from sunflower leaves by an enzymic procedure. The cell suspensions possessed high photosynthesis rates. The products of cell photosynthesis were similar to the products of leaf disc photosynthesis. The relatively high radioactivity incorporated into malate after ¹⁴CO₂ feeding suggests that PEP carboxylase might participate in CO₂ fixation. Sunflower leaf extracts possessed a PEP carboxylase activity slightly higher than that of other C₃ species. Inhibition of PEP carboxylase by maleate decreased cell photosynthesis by only 15% and the first products of cell photosynthesis were phosphorylated compounds. It is concluded that the high photosynthesis rates displayed by sunflower are not due to a parallel C₄ pathway of photosynthesis but are rather dependent, at least in part, on the activity, or the amount, of RuBP carboxylase.Abbreviations PVP polyvinylpyrrolidone - PDS potassium dextran sulfate - DTT dithiothreitol - PEG polyethyleneglycol - RuBP ribulose 1,5-bisphosphate - PEP phosphoenolpyruvate - Mes 2-(N-morpholino) ethanesulfonic acid - Hepes N-2-hydroxyethylpiperazine-N-2-ethanesulfonic acid     

Geographical and environmental distribution of C3 and C4 grasses in the Sinai,Negev, and Judean deserts

Summary The relation between photosynthetic pathway and habitat of the grass species recorded in the desert regions of Sinai, Negev, and Judea was investigated. The climatic conditions and micro-environments in the study area vary considerably, and the distribution of the various species is found to conform to specific patterns which reveal the adaptive advantages of the different photosynthetic pathways. There is also a distinct correlation between the phytogeographic origin of the grass species and the photosynthetic pathways that they utilize.The survey shows that the majority of the grass species in the region are of the C₃ type and all except one of these species belong to the Holarctic domain. This is in accordance with the fact that the region forms part of the Mediterranean winter rainfall regime and that C₃ species have an adaptive advantage where minimum temperatures are low during the winter growing season.The occurence of C₄ species increases with decreasing rainfall and they dominate in those districts where temperatures are high throughout the year. These C₄ grasses are of both Holarctic and Palaeotropic origin according to the classification adopted here, but they are essentially all elements of the Saharo-Arabian, Irano-Turanian, Sudanian, or Tropical phytogeographic regions and are not typical of the Mediterranean or Euro-Siberian floras. The plants with multi-regional distributions that occur in Mediterranean communities may well be intrusive.Analysis of the three subtypes of the C₄ species suggests that the malate-forming NADP-me grasses grow where water stress is not a dominating factor, while the aspartateforming NAD-me grasses are more successful under xeric conditions. The PEP-ck species are not abundant and form an intermediate group between the NADP-me and NAD-me subtypes.     

A comparative immunocytochemical localization study of phosphoenolpyruvate carboxylase in leaves of higher plants

The intracellular localization of phosphoenolpyruvate (PEP) carboxylase in plants belonging to the C₄, Crassulacean acid metabolism (CAM) and C₃ types was invetigated using an immunocytochemical method with an immune serum raised against the sorghum leaf enzyme. The plants studied were sorghum, maize (C₄ type), kalanchoe (CAM type), french bean, and spinach (C₃ type). In the green leaves of C₄ plants, it was shown that the carboxylase was located in the mesophyll and stomatic cells, being largely cytosolic in the mesophyll cells. Similarly, in CAM plants, the enzyme was found mainly outside the chloroplasts. In contrast, in C₃ plants, the PEP carboxylase appeared to be distributed between the cytosol and the chloroplasts of foliar parenchyma. Examination of sections from etiolated leaves showed fluorescence emission from etioplasts and cytosol for the parenchyma of french bean as well as for the bundle sheath and mesophyll of sorghum leaves. This data indicated that during the greening process photoregulation and evolution of PEP carboxylase is dependent on the tissue and on the metabolic type of the plant considered.Abbreviations CAM Crassulacean acid metabolism - PEP phosphoenolpyruvate     

Carbon-isotope discrimination by leaves of Flaveria species exhibiting different amounts of C3-and C4-cycle co-function

Carbon-isotope ratios were examined as ¹³C values in several C₃, C₄, and C₃–C₄ Flaveria species, and compared to predicted ¹³C, values generated from theoretical models. The measured ¹³C values were within 4 of those predicted from the models. The models were used to identify factors that contribute to C₃-like ¹³C values in C₃–C₄ species that exhibit considerable C₄-cycle activity. Two of the factors contributing to C₃-like ¹³C values are high CO₂ leakiness from the C₄ pathway and pi/pa values that were higher than C₄ congeners. A marked break occurred in the relationship between the percentage of atmospheric CO₂ assimilated through the C₄ cycle and the ¹³C value. Below 50% C₄-cycle assimialtion there was no significant relationship between the variables, but above 50% the ¹³C values became less negative. These results demonstrate that the level of C₄-cycle expression can increase from, 0 to 50% with little integration of carbon transfer from the C₄ to the C₃ cycle. As expression increaces above 50%, however, increased integration of C₃- and C₄-cycle co-function occurs.Abbreviations and symbols RuBP carboxylase ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) - PEP carboxylase phosphoenolpyruvate carboxylase (EC 4.1.1.31) - pa atmospheric CO₂ partial pressure - pi intercellular CO₂ partial pressure - isotope ratio - quantum yield for CO₂ uptake     

The quantum yield for CO2 uptake in C3 and C4 grasses

The quantum yield for CO₂ uptake was measured in C₃ and C₄ monocot species from several different grassland habitats. When the quantum yield was measured in the presence of 21% O₂ and 340 cm³ m^-3 CO₂, values were very similar in C₃ monocots, C₃ dicots, and C₄ monocots (0.045–0.056 mole CO₂ · mole^-1 quanta absorbed). In the presence of 2% O₂ and 800 cm³ m^-3 CO₂, enhancements of the quantum yield values occurred for the C₃ plants (both monocots and dicots), but not for C₄ monocots. A dependence of the quantum yield on leaf temperature was observed in the C₃ grass, Agropyron smithii, but not in the C₄ grass, Bouteloua gracilis, in 21% O₂ and 340 cm³ m^-3 CO₂. At leaf temperatures between 22–25°C the quantum yield values were approximately equal in the two species.