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。     

不同土壤水分条件下土壤容重对玉米木质部汁液中ABA浓度和气孔导度的影响

受旱玉米植株木质部汁液中的ABA浓度是高水分处理的5倍,土壤容重每增加0.12g·cm-3,木质部液汁中ABA浓度约增加1倍。在相同土壤基质势下,植株的气孔导度和蒸腾速率随土壤容重的增大而下降,而容重对植株的叶水势没有影响。生长在混合容重(一边为1.20g·cm-3,另一边为1.45g·cm-3)土壤上的植株中,ABA浓度和气孔导度与全部根系处在高容重土壤中的植株接近。     

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。     

不同R∶FR值对菊花叶片气孔特征和气孔导度的影响

以切花菊品种"神马(Jinba)"为试材,2010年10月至2011年2月间在南京信息工程大学试验温室采用不同Red (660±10) nm: Far-red (730±10) nm值的LED光源短日处理,研究了温室切花菊叶片气孔特征和气孔导度对不同R∶FR值的响应。结果表明:不同R∶FR值短日处理35d菊花叶片的上表皮和下表皮的气孔直径分别以R∶FR值4.5和6.5处理最大,均以R∶FR值2.5处理最小,气孔密度和气孔开度均以R∶FR=2.5处理最高,以R∶FR值6.5处理最低,下表皮的气孔密度、气孔开度明显高于上表皮;不同R∶FR值处理叶片的气孔开张比和气孔指数差异不显著;在相同光强下,菊花叶片气孔导度和光合速率由大到小的R∶FR值顺序依次为2.5、4.5、0.5、6.5。叶片气孔导度与气孔指数、气孔密度、气孔开张比和气孔开度成正相关,与气孔长度和气孔宽度呈负相关;R∶FR值在2.5-6.5范围内,随光质中红光成分增加,叶片气孔密度、气孔指数、气孔开度、气孔开张比和气孔导度显著降低。     

分根区干湿交替对玉米光合速率及蒸腾效率的影响

以玉米为材料研究了分根区干湿交替对叶片光合速度及蒸腾效率的影响,结果发现控制１／２根区交替供水和固定１／２根区供水在减少用水量的同时,叶片蒸腾速度明显下降,而光合速度不明显,蒸腾效率提高;控制１／２根区交替供水比固定１／２根区供水在光合维持高水平稳定时所要求的土壤含水量低,其蒸腾效率较高。干湿交替能够增加根冠比、根干重和根九。证明控制性分根交灌溉可提高玉米腾效率达到节水的目的。     

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。     

春玉米持续干旱过程中常用气孔导度模型的比较研究

气候变化背景下,干旱频发导致的土壤水分变化将影响气孔导度模型的适用性,进而影响生态系统碳-氮-水循环模拟的准确性。基于春玉米持续干旱田间模拟试验资料,比较了常用气孔导度模型在干旱条件下的模拟效果,评价了土壤水分响应函数对气孔导度模型效果的影响,并探讨了气孔导度模型的适用土壤水分范围。结果表明,在持续干旱过程中,模型模拟效果表现为BBL模型最优,其次是USO模型和BWB模型,Jarvis模型最差;引入土壤水分响应函数,提高了BWB模型和USO模型的模拟效果,而降低了Jarvis模型和BBL模型模拟效果,模型模拟效果表现为USO修正模型最优,其次是BBL修正模型和BWB修正模型,Jarvis修正模型最差。在持续干旱过程中,Jarvis模型和BWB模型的剩余气孔导度较大,而BBL模型和USO模型的剩余气孔导度相对较小,表明BBL模型和USO模型在干旱条件下具有一定的稳定性。基于95%置信区间判断表明:Jarvis模型、BBL模型和USO模型在土壤相对湿度范围为33%—83%条件下适用,而BWB模型的适用土壤相对湿度范围为33%—76%,引入水分响应函数后可在试验条件下适用。研究结果可为干旱条件下选取合适的气孔导度模型以准确模拟陆地生态系统碳循环和水循环提供依据,并为改善农业水资源的有效使用和评估提供支撑。     

干旱条件下ABA与气孔导度和叶片生长的关系

介绍了干旱情况下ＡＢＡ的产生、运输,以及根源ＡＢＡ与气孔导度和叶片生长之间的定性、定量关系,并对干旱时叶片ＡＢＡ的重新分布,以及引起气孔关闭和抑制生长的机理作了叙述。     

黄土丘陵区沙棘气孔导度及其影响因子

依据 1 998年在半干旱黄土丘陵区安塞的观测资料 ,对沙棘适应该地区生境的气孔运动进行了分析研究。结果表明 :(1 )沙棘气孔导度日均值为 2 1 4.47mmol· m- 2 · s- 1,日变化曲线呈双峰型。 (2 )沙棘气孔导度与 CO2 浓度间有十分显著的相关关系 ,复相关系数为0 .96 3 5 ;与光合有效辐射、气温、叶水势间有显著的相关关系 ,复相关系数为 0 .70 3 3～0 .8971 ;但与相对湿度间相关性不显著。 (3 )沙棘适应半干旱黄土丘陵区生境的气孔调节机制为反馈式反应 ,即由于叶水势降低导致气孔导度减小 ,从而减少蒸腾耗水 ,达到节约用水 ,适应干旱的目的 ;(4)沙棘的水分利用效率随气孔限制值的增大而减小 ,二者间呈显著的负相关。这为了解沙棘的抗旱机理 ,选育节水抗旱品种 ,提高水分利用效率提供了科学依据     

Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

Mesophyll conductance to CO2 and Rubisco as targets for improving intrinsic water use efficiency in C3 plants

Water limitation is a major global constraint for plant productivity that is likely to be exacerbated by climate change. Hence, improving plant water use efficiency (WUE) has become a major goal for the near future. At the leaf level, WUE is the ratio between photosynthesis and transpiration. Maintaining high photosynthesis under water stress, while improving WUE requires either increasing mesophyll conductance (g_m) and/or improving the biochemical capacity for CO₂ assimilation—in which Rubisco properties play a key role, especially in C₃ plants at current atmospheric CO₂. The goals of the present analysis are: (1) to summarize the evidence that improving g_m and/or Rubisco can result in increased WUE; (2) to review the degree of success of early attempts to genetically manipulate g_m or Rubisco; (3) to analyse how g_m, g_sw and the Rubisco's maximum velocity (V_cmax) co‐vary across different plant species in well‐watered and drought‐stressed conditions; (4) to examine how these variations cause differences in WUE and what is the overall extent of variation in individual determinants of WUE; and finally, (5) to use simulation analysis to provide a theoretical framework for the possible control of WUE by g_m and Rubisco catalytic constants vis‐à‐vis g_sw under water limitations.     

Xylem ABA controls the stomatal conductance of field-grown maize subjected to soil compaction or soil drying

Stomatal conductance of individual leaves was measured in a maize field, together with leaf water potential, leaf turgor, xylem ABA concentration and leaf ABA concentration in the same leaves. Stomatal conductance showed a tight relationship with xylem ABA, but not with the current leaf water status or with the concentration of ABA in the bulk leaf. The relationship between stomatal conductance and xylem [ABA] was common for variations in xylem [ABA] linked to the decline with time of the soil water reserve, to simultaneous differences between plants grown on compacted, non-compacted and irrigated soil, and to plant-to-plant variability. Therefore, this relationship is unlikely to be fortuitous or due to synchronous variations. These results suggest that increased concentration of ABA in the xylem sap in response to stress can control the gas exchange of plants under field conditions.     

Ontogenetic changes in stomatal size and conductance of sunflowers

The ontogenetic changes in stomatal size, frequency and conductance (g_s) on abaxial and adaxial leaf surfaces of sunflower plants (Helianthus annuus L. Russian Mammoth) were examined under controlled environmental conditions. The stomatal frequency on the adaxial and abaxial leaf surfaces decreased with leaf ontogeny and insertion level. The ratio of adaxial to abaxial stomatal frequency did not change with leaf ontogeny and insertion level, and 42–44% of total stomata was apportioned to the adaxial surface. Ontogenetic changes in stomatal pore length were detected and increased with ontogenesis. The stomatal length of both leaf surfaces had linear relationships with leaf area. Ontogenetic changes in g_s were similar between the two surfaces. However the adaxial g_s was lower than abaxial g_s in leaves of higher insertion levels. Conductance had a linear relationship with width x frequency but not with pore area.     

Use of thermography for quantitative studies of spatial and temporal variation of stomatal conductance over leaf surfaces

This paper describes a new approach to the calibration of thermal infrared measurements of leaf temperature for the estimation of stomatal conductance and illustrates its application to thermal imaging of plant leaves. The approach is based on a simple reformulation of the leaf energy balance equation that makes use of temperature measurements on reference surfaces of known conductance to water vapour. The use of reference surfaces is an alternative to the accurate measurement of all components of the leaf energy balance and is of potentially wide application in studies of stomatal behaviour. The resolution of the technique when applied to thermal images is evaluated and some results of using the approach in the laboratory for the study of stomatal behaviour in leaves of Phaseolus vulgaris L. are presented. Conductances calculated from infrared measurements were well correlated with estimates obtained using a diffusion porometer.     

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。     

玉米叶片水分利用效率的保守性

水分利用效率是植物个体或生态系统水分利用过程的重要特征参数,可表征不同时空尺度的植物碳-水耦合关系,对植物适应气候变化研究具有重要意义。以玉米为例,利用中国气象局固城农业气象野外科学试验基地2013—2014年玉米不同灌溉方案模拟试验资料,对不同叶位叶片的水分利用效率特征及其影响因素进行分析。结果表明:植株顶部第1片叶片水分利用效率在拔节期和乳熟期呈现明显的峰值,反映出明显的周期变化规律及其与叶片生理生态特征的紧密相关。在相同环境条件下,不同叶位叶片的水分利用效率不存在显著性差异,即玉米叶片水分利用效率具有空间稳定性与叶龄保守性。同时,研究指出叶片光合速率和蒸腾速率在叶位之间的协调变化是导致空间稳定性和叶龄保守性的主要原因。研究结果可为植物水分关系研究提供参考,也可为水分利用效率的尺度化研究提供依据。     

High yields in advanced lines of Pima cotton are associated with higher stomatal conductance, reduced leaf area and lower leaf temperature

Advanced lines of Pima cotton ( Gossypium barbadense L.) bred for higher yield potential and heat resistance have higher stomata conductance and smaller leaf areas than those of obsolete lines. In controlled experiments, five commercial lines of Pima cotton having increasing lint yield and heat resistance showed a gradient of increasing stomatal conductance and decreasing leaf size. In field experiments, heat-sensitive, low yield Pima lines showed a lower stomatal conductance than high yielding, advanced lines. This indicates that selection for high yield potential and heat resistance has imposed a selection pressure for higher stomatal conductance and smaller leaf areas. The higher stomatal conductance and smaller leaf area in the advanced lines resulted in a lower leaf temperature in both controlled environments and in the field. The largest leaf temperature differences between obsolete and advanced lines were observed in the afternoon. These differences coincided with the largest differences in stomatal conductance and the highest air temperatures. Measurements of stomatal conductance and leaf temperature in field-grown progeny from a cross between the advanced line, Pima S-6. and the obsolete line, Pima 32, showed that genetically determined differences in stomatal conductance resulted in corresponding differences in leaf temperature. None of the altered physiological traits were selected for in the breeding program, indicating that selection for the desired agronomic traits imposed selection pressures on the altered physiological traits. The increases in stomatal conductance and decreases in leaf area could represent an integrated response to selection pressures on enhanced evaporative cooling, ensuing from selection for heat resistance.