冬小麦叶片气孔导度模型水分响应函数的参数化

植物气孔导度模型的水分响应函数用来模拟水分胁迫对气孔导度的影响过程, 是模拟缺水环境下植物与大气间水、碳交换过程的关键算法。水分响应函数包括空气湿度响应函数和土壤湿度(或植物水势)响应函数, 该研究基于田间实验观测, 分析了冬小麦(Triticum aestivum)叶片气孔导度对不同空气饱和差和不同土壤体积含水量或叶水势的响应规律。一个土壤水分梯度的田间处理在中国科学院禹城综合试验站实施, 不同水分胁迫下的冬小麦叶片气体交换过程和气孔导度以及其他的温湿度数据被观测, 同时观测了土壤含水量和叶水势。实验数据表明, 冬小麦叶片气孔导度对空气饱和差的响应呈现双曲线规律, 变化趋势显示大约1 kPa空气饱和差是一个有用的阈值, 在小于1 kPa时, 冬小麦气孔导度对空气饱和差变化反应敏感, 而大于1 kPa后则反应缓慢; 分析土壤体积含水量与中午叶片气孔导度的关系发现, 中午叶片气孔导度随土壤含水量增加大致呈现线性增加趋势, 但在平均土壤体积含水量大于大约25%以后, 气孔导度不再明显增加, 而是维持在较高导度值上下波动; 冬小麦中午叶片水势与相应的气孔导度之间, 随着叶水势的增加, 气孔导度呈现增加趋势。根据冬小麦气孔导度对空气湿度、土壤湿度和叶水势的响应规律, 研究分别采用双曲线和幂指数形式拟合了水汽响应函数, 用三段线性方程拟合了土壤湿度响应函数和植物水势响应函数, 得到的参数可以为模型模拟冬小麦的各类水、热、碳交换过程采用。     

植物气孔导度的机理模型

Ball-Berry气孔导度模型及其修正模型是评价植物叶片气孔调节的重要工具。该文从CO₂分子在叶片气孔中扩散这个最基本的物理过程出发, 应用物理学中的分子扩散和碰撞理论、流体力学与植物生理学等知识, 严格推导出叶片气孔导度的机理模型。利用美国Li-Cor公司生产的Li-6400光合作用测定仪控制CO₂浓度、湿度和温度, 测量了华北平原冬小麦(Triticum aestivum)的光响应数据和气孔导度数据。拟合结果表明: 推导的气孔导度机理模型较之Ball-Berry气孔导度模型和Tuzet等气孔导度模型, 能更好地描述冬小麦的气孔导度与净光合速率之间的关系。如果用气孔导度的机理模型耦合光合作用对光响应的修正模型, 则耦合模型可以很好地描述华北平原冬小麦叶片气孔导度对光强的响应曲线, 并可直接估算冬小麦的最大气孔导度和对应的饱和光强, 同时可以研究最大气孔导度是否与最大净光合速率同步的问题。拟合结果还表明: 冬小麦在30 ℃、560 μmol·mol^-1CO₂, 或在32 ℃、370 μmol·mol^-1CO₂条件下, 最大气孔导度与最大净光合速率并不同步。     

羊草气孔导度的Jarvis-类模型

在干旱半干旱气候条件下,土壤水分状况通常是决定植物气孔导度的重要因素,现有气孔导度模型Jarvis-类和耦合模型(或光合-导度模型)未充分考虑这一因素对气孔导度的影响。本文以Jarvis气孔导度模型为基础,提出一个充分考虑土壤水分状况因素的气孔导度模型。该模型对羊草连续两年(1998~1999)野外实地观测结果拟合良好(R2=0.603),预测能力较线性回归方程(R2=0.361)有明显提高。     

冬小麦群体叶片气孔导度差异性分析

对冬小麦 (Triticum aestivum L.)两个品种 3个生育期测定和分析了群体叶片的气孔导度。结果表明 ,近轴面气孔导度比远轴面的大 ,二者之比呈近似正态分布 ,最大概率出现约为 1 .5,该比值在冠层垂直方向变化不显著 ,但随季节而变 ,抽穗期最大。叶片气孔导度从冠层顶部向下迅速递减 ,递减系数约为 0 .57,且各层日变化大致平行。冠层上部气孔活跃 ,导度日变化明显 ,下部日变化不大。同一叶片的气孔导度由近轴面的叶基 ,经由其叶中、叶尖 ,翻过尖点 ,再从远轴面的叶尖经叶中向叶基递减。对近轴面叶中、叶尖气孔导度与该面平均气孔导度相关系数较大 ,而对远轴面叶中、叶基气孔导度与该面气孔导度相关系数较大 ,据此建议测定近轴面气孔导度时 ,应以叶中、叶尖部位为最佳点 ,而测定远轴面气孔导度时 ,应以叶中、叶基部位为最佳点     

最优气孔行为理论和气孔导度模拟

气孔调节功能是陆地生态系统碳-水耦合过程中最重要的环节。与即时的气孔导度测量相比, 气孔导度斜率能有效地反映气孔导度对CO₂浓度、饱和水汽压亏缺和光合作用的敏感性, 包含了环境因子对光合作用和临界水分利用效率等的综合影响, 为研究全球变化下陆地生态系统碳-水耦合关系提供了一个简明且综合的框架。气孔导度模型从经验模型、半经验模型发展到机理模型, 经过很多学者的改进, 但是模型参数的生物学意义和变化规律还不明确。鉴于气孔导度斜率方面研究的重要性和研究的不足, 为了加强对气孔导度调节规律的认识, 并减少气孔导度模拟的不确定性, 该文主要综述了长期以来国内外关于最优气孔行为理论和气孔导度模拟方面的研究成果, 其中包括广泛使用的气孔导度模型及参数意义, 讨论影响气孔导度斜率的主要因素以及气孔导度机理模型的应用, 并对最优气孔行为理论和气孔导度模拟方面的研究做了简单展望。     

羊草叶片气孔导度对环境因子的响应模拟

准确定量描述植物气孔对环境的响应是了解植物光合作用机理、预测植物生产力及其大气－植被－土壤系统中水分和热量交换的关键。利用松嫩平原盐碱化草地羊草光合生理特征的野外观测数据,分析了羊草叶片气孔导度对环境因子的反应,结果表明：羊草叶片气孔导度对环境因子变化敏感,尤其对瞬时光合有效辐射（ＰＡＲ）、叶片与空气间的水汽压亏损（ＶＰＤ）和空气温度（Ｔａ）反应十分明显。依据野外实测资料对国际上两类代表性气孔导度     

羊草叶片气孔导度特征及数值模拟

对松嫩平原草地羊草叶片气孔导特征及与环境因子关系的研究结果表明,羊草叶片气孔导度日变化与环境因子密切相关,晴天表现为双峰曲线,阴天为单峰曲线,同时叶片气孔导度（gs)对瞬时光合有效辐射（PAR）,叶片与空气间的水汽压亏损（VPD）,空气温度（Ta）反应十分明显,依据野外实测资料,在对国际上两类代表性气孔导度模型验证比较的基础上,建立了适用于羊草草原的羊草叶片气孔导度对环境因子的响应模型gs=PAR(2.01Ta^2 147.74Ta-2321.11)/(444.62 PAR)(-538.04 VPD).     

基于叶片尺度观测数据的气孔导度模型评价

气孔导度(g)是控制冠层与大气之间能量和水分交换的重要因素。空气湿度是控制植物叶片气孔导度的一个关键环境因子。在过去的十几年中,普遍得到应用的是Ball-Woodrow-Berry(BWB)模型和Leuning模型中气孔导度与湿度的关系。本研究使用一个诊断变量f(H),基于农田叶片水平的光合-气孔导度观测数据,对BWB模型、Leuning模型以及新发展的power-h模型和power-D模型进行了气孔导度模拟效果的比较和评价。结果表明:BWB模型描述的是g和相对湿度(hs)之间的一种线性关系,当空气较为湿润时,模拟结果存在较大的低估;Leuning模型中反映的是g与饱和水汽压差(Ds)的非线性函数,降低了模拟结果的误差,但仍然不能很好地描述g在较湿状况下的显著升高;相比之下,两个新的模型,即Ds的指数函数和(1-hs)的指数函数形式模型能提高模拟结果的精度。这个研究结果也表明基于Ds的模型模拟效果要好于基于hs的模型。     

冬小麦气孔臭氧通量拟合及通量产量关系的比较分析

基于田间原位开顶箱(Open-Top Chambers,OTCs)实验,研究了不同浓度臭氧(O3)处理下(自然大气处理,AA;箱内大气处理,NF;箱内低浓度O3处理,NF+40 nL/L;箱内中等浓度O3处理,NF+80 nL/L;箱内高浓度O3处理,NF+120 nL/L),冬小麦(Triticum aestivum L.)旗叶气孔运动对不同环境因子的响应,并通过剂量反应分析,比较了冬小麦产量损失与累积气孔O3吸收通量(AFstX)和累积O3暴露浓度(AOT40和SUM06)的相关性差异。结果表明:冬小麦旗叶气孔运动的光饱和点和最适温度分别约为400μmol.m-.2s-1和27℃,水汽压差、土壤含水量和O3剂量的气孔限制临界值分别约为1.4 kPa、-100 kPa和20μL.L-.1h-1,超过此临界值时,气孔导度会明显下降。利用Jarvis气孔导度模型对冬小麦旗叶气孔导度和气孔O3吸收通量进行了预测,结果表明Jarvis模型解释了冬小麦实测气孔导度60%的变异性。由于不同时期植物体气孔导度的差异,冬小麦旗叶生长期内累积气孔O3吸收通量(AFstX)呈非线性增加趋势。O3吸收速率临界值(X)为4 nmol.m-.2s-1时,累积O3吸收通量(AFst4)与冬小麦产量的相关性最高(R2=0.76),该数值介于O3暴露指标AOT40和SUM06的剂量反应决定系数(0.74和0.81)之间。与O3浓度指标(AOT40和SUM06)相比,O3通量指标(AFstX)在本试验冬小麦产量损失评价中未表现出明显优势。     

麦田冠层气孔导度的分层研究

小麦灌浆期和乳熟期冠层各层叶片上、下表面的气孔导度之间呈正相关关系;冠层不同层的叶片气孔导度从早到傍晚有平行变化的趋势,数值上存在较大的差异,一般从冠层上到下递减。经分析,这主要与冠层叶片接受的光强自上而下递减有关,且这时所对应的叶片水势自冠层上到下递增的幅度大。测算结果表明,冠层气孔导度白天亦呈明显的日变化,灌浆期的值大于乳熟期的值。     

Characteristics and partitioning of ozone dry deposition measured by eddy-covariance technology in a winter wheat field

Aims Anthropogenic pollutants cause an increase in ground-level ozone concentration, which is a known threat to plant growth and yield and has been extensively observed worldwide. Since ozone is only slightly soluble in water, it is deposited mainly through dry deposition in terrestrial ecosystem. The object of this study was to analyze the characteristics of ozone dry deposition and to estimate the contribution of stomatal and non-stomatal ozone deposition pathways to total ozone deposition in a winter wheat field.Methods The research site was a winter wheat (Triticum aestivum) field located in Yongfeng experimental station of Nanjing University of Information Science & Technology. The data used in this study were collected from March 16, 2016 to May 30, 2016. We observed ozone dry deposition with an eddy-covariance system. This system mainly included a 3D sonic anemometer, an open-path infrared absorption spectrometer, a fast-response ozone chemiluminescent analyzer and a slow-response ozone monitor. We simultaneously measured meteorological data including solar radiation (SR), air temperature (T), air relativity humidity (RH), wind speed, net radiation, and rainfall. All raw data were recorded with data-logger and averaged every 30 min.Important findings Half hourly means of ozone concentrations (C_O3), ozone flux (F_O3) and ozone dry deposition velocity (V_d) in the winter wheat field were 32.9 nL·L^-1, -5.09 nmol·m^-2·s^-1, 0.39 cm·s^-1, and the ranges of them were 16-58 nL·L^-1, -2.9- -11.7 nmol·m^-2·s^-1, 0.17-0.63 cm·s^-1, respectively. F_O3 and C_O3/V_d were found to be mismatched with phase peaks occurring at different time intervals. The ecosystem was more effective on ozone dry deposition, under conditions of moderate to high SR (SR ≥ 400 W·m^-2), moderate T and humility (T = 18 °C and RH > 40%). The relationship between V_dmax and SR was this function (y = 1.06 -exp (-0.0094 - x)). V_dmax increased with SR When SR < 400 W·m^-2, and V_dmax reached its maximum when SR =400 W·m^-2. V_dmax maintained its maximum when SR ≥ 400 W·m^-2. The relationship between V_dmax and T was “bell” curve (y = 1.06 - (x - 18)²/169). V_dmax reached its maximum when T = 18 °C. V_dmax decreased with RH when RH < 40 % (y = 0.030x - 0.106). The variation of V_d might uncertainty when RH was high. There was a liner positive relationship between friction velocity (u*) and V_d, but this relationship was not significant. The mean day-to-day and daytime contributions of stomatal and non-stomatal ozone deposition pathway to total ozone deposition were 32%, 68% and 42%, 58%, respectively, during the whole experimental period.     

Suitability of a combined stomatal conductance and photosynthesis model for calculation of leaf-level ozone fluxes

Currently, the most important source of uncertainty in stomatal ozone flux ( FO3) modelling is the stomatal conductance ( gst) factor. Hence FO3 model accuracy will strongly depend on the gst model being implemented. In this study the recently developed semi-empirical Gst model of Dewar was coupled to the widely known biochemical photosynthesis ( An) model of Farquhar. The Gst performance of this model combination was evaluated with a 4-month time series of beech ( Fagus sylvatica L.) measurements. The Gst model was hereto optimized in two steps to a 4-day and a 8-day period. A comparison between the modelled and measured gst to O(3) (gstO3) revealed a rather good overall performance (R(2)=0.77). Errors between the model combination and the measurements are thought to be largely caused by a moderate performance of the AN model, due to poor parameterization. Two 2-day periods with distinctly differing soil and meteorological conditions were chosen to give a picture of the daily gst performance. Although instant relative differences between modelled and measured gstO3 are sometimes high, the model combination is able to simulate the rough daily courses of gstO3 and hence FO3 reasonably well. Further improvement on full parameterization of the gst model and a well-parameterized An model to be linked to are needed to draw founded conclusions about its performance. Future efforts hereto are certainly justified since the model's mechanistic nature makes it a tool able to model gst variation in space and time, O(3) effects on gst, and effective FO3.     

陆地生态系统臭氧通量观测和气孔吸收估算研究进展

近地面大气中臭氧(O3)对植物生长发育和产量会产生不良影响。工业和交通排放的增加使得全球地面O3浓度逐年增加,不断升高的O3浓度已开始影响到我国的粮食产量。O3对植物的影响是由于其进入植物体内发生生化反应所引起的,所以需要建立一种考虑到植物生理生态状况的评估指标来评估O3对植物的影响。其中基于O3通量(特别是植物气孔吸收)的评价指标和方法,被认为比传统的基于O3浓度的评价指标和方法更符合O3对植物的影响机理。介绍了O3对生态系统影响评估方法和评价指标,重点评述了生态系统尺度O3通量观测和气孔吸收估算的主要方法以及在不同生态系统上的研究进展分析了我国关于O3对植物和生态系统影响的研究现状,并对未来的研究工作进行了展望。     

Responses of stomatal conductance to light, humidity and temperature in winter wheat and barley grown at three concentrations of carbon dioxide in the field

Responses of leaf stomatal conductance to light, humidity and temperature were characterized for winter wheat and barely grown at ambient (about 350 μmol mol^?1 in the daytime), ambient + 175 and ambient + 350 μmol mol^?1 concentrations of carbon dioxide in open‐topped chambers in field plots over a three year period. Stomatal responses to environment were determined by direct manipulation of single environmental factors, and those results were compared with responses derived from natural day to day variation in mid‐day stomatal conductance. The purpose of these experiments was to determine the magnitude of reduction in stomatal conductance at elevated [CO₂], and to assess whether the relative response of conductance to elevated [CO₂] was constant across light, humidity and temperature conditions. The results indicated that light, humidity and temperature all significantly affected the relative decrease in stomatal conductance at elevated [CO₂]. The relative decrease in conductance with elevated [CO₂] was greater at low light, low water vapour pressure difference, and high temperature in both species. For measurements made at saturating light near mid‐day, the ratio of mid‐day stomatal conductances at doubled [CO₂] to that at ambient [CO₂] ranged from 0.42 to 0.86, with a mean of 0.66 in barley, and from 0.33 to 0.80, with a mean of 0.56 in wheat. Day‐to‐day variation in the relative effect of elevated [CO₂] on conductance was correlated with the relative stimulation of [CO₂] assimilation rate and with temperature. Some limitations of multiple linear regression, multiplicative, and ‘Ball–Berry' models as summaries of the data are discussed. In barley, a better fit to the models occurred in individual years than for the combined data, and in wheat a better fit to the models occurred when data from near the end of the season were removed.     

冬麦田臭氧干沉降过程的观测

地表臭氧作为近地层最主要的大气污染物之一,其不断上升的浓度及其对粮食作物的影响受到越来越多的关注.本研究利用微气象学观测方法,探明自然条件下冬麦田的臭氧沉降过程,分析了影响臭氧沉降过程的环境因子.结果表明: 观测期内(小麦生长旺期)臭氧通量均值为-0.35 μg·m^-2·s^-1(负号表示沉降方向指向地面),臭氧沉降平均速率为0.55 cm·s^-1,空气动力学阻力均值为30 s·m^-1,粘性副层阻力均值为257 s·m^-1,冠层阻力均值为163 s·m^-1,且均存在明显的日变化趋势.臭氧沉降阻力大小受摩擦速度、太阳辐射强度、温度和相对湿度等因素的影响.     

臭氧胁迫下冬小麦物质生产与分配的数值模拟

利用ML9810B型臭氧监测分析仪,测定了浙江嘉兴麦田空气O3浓度,并通过改进的开顶式气室实验确定O3浓度变化对冬小麦叶片光合速率的影响函数.在此基础上,加入O3对叶片生长和穗部光合影响的模拟函数,建立反映O3对冬小麦生长和产量形成影响的作物模型.模型的检验结果表明,该模型较好地反映了O3对冬小麦生长的影响,生物量平均相对误差为10.3%.对冬小麦春后生育期(3—5月)的研究表明,水肥适宜时,由O3影响造成的该地区冬小麦干物质累积总损失量为11.4%,产量损失为17.8%.     

Ozone‐induced stomatal sluggishness changes stomatal parameters of Jarvis‐type model in white birch and deciduous oak

• Stomatal ozone flux is closely related to ozone injury to plants. Jarvis‐type multiplicative model has been recommended for estimating stomatal ozone flux in forest trees. Ozone can change stomatal conductance by both stomatal closure and less efficient stomatal control (stomatal sluggishness). However, current Jarvis‐type models do not account for these ozone effects on stomatal conductance in forest trees.
• We examined seasonal course of stomatal conductance in two common deciduous tree species native to northern Japan (white birch: Betula platyphylla var. japonica ; deciduous oak: Quercus mongolica var. crispula ) grown under free‐air ozone exposure. We innovatively considered stomatal sluggishness in the Jarvis‐type model using a simple parameter, s , relating to cumulative ozone uptake (defined as POD : phytotoxic ozone dose).
• We found that ozone decreased stomatal conductance of white birch leaves after full expansion (?28%). However, such a reduction of stomatal conductance by ozone fell in late summer (?10%). At the same time, ozone reduced stomatal sensitivity of white birch to VPD and increased stomatal conductance under low light conditions. In contrast, in deciduous oak, ozone did not clearly change the model parameters.
• The consideration of both ozone‐induced stomatal closure and stomatal sluggishness improved the model performance to estimate stomatal conductance and to explain the dose–response relationship on ozone‐induced decline of photosynthesis of white birch. Our results indicate that ozone effects on stomatal conductance (i.e . stomatal closure and stomatal sluggishness) are crucial for modelling studies to determine stomatal response in deciduous trees, especially in species sensitive to ozone.

     

Genotypic differences in leaf biochemical, physiological and growth responses to ozone in 20 winter wheat cultivars released over the past 60 years

Ozone (O₃) concentrations in periurban areas in East Asia are sufficiently high to decrease crop yield. However, little is known about the genotypic differences in O₃ sensitivity in winter wheat in relation to year of cultivar release. This paper reports genotypic variations in O₃ sensitivity in 20 winter wheat cultivars released over the past 60 years in China highlighting O₃‐induced mechanisms. Wheat plants were exposed to elevated O₃ (82 ppb O₃, 7 h day^?1) or charcoal‐filtered air (<5 ppb O₃) for 21 days in open top chambers. Responses to O₃ were assessed by the levels of antioxidative activities, protein alteration, membrane lipid peroxidation, gas exchange, leaf chlorophyll, dark respiration and growth. We found that O₃ significantly reduced foliar ascorbate (?14%) and soluble protein (?22%), but increased peroxidase activity (+46%) and malondialdehyde (+38%). Elevated O₃ depressed light saturated net photosynthetic rate (?24%), stomatal conductance (?8%) and total chlorophyll (?11%), while stimulated dark respiration (+28%) and intercellular CO₂ concentration (+39%). O₃ also reduced overall plant growth, but to a greater extent in root (?32%) than in shoot (?17%) biomass. There was significant genotypic variation in potential sensitivity to O₃ that did not correlate to observed O₃ tolerance. Sensitivity to O₃ in cultivars of winter wheat progressed with year of release and correlated with stomatal conductance and dark respiration in O₃‐exposed plants. O₃‐induced loss in photosynthetic rate was attributed primarily to impaired activity of mesophyll cells and loss of integrity of cellular membrane as evidenced by increased intercellular CO₂ concentration and lipid peroxidation. Our findings demonstrated that higher sensitivity to O₃ in the more recently released cultivars was induced by higher stomatal conductance, larger reduction in antioxidative capacity and lower levels of dark respiration leading to higher oxidative damage to proteins and integrity of cellular membranes.