姜寒冰^{1, 2,},
张玉翠¹,
任晓东^{1, 3},
要家威^{1, 2},
沈彦俊^1,,
1.中国科学院农业水资源重点实验室/河北省节水农业重点实验室/中国科学院遗传与发育生物学研究所农业资源研究中心 石家庄 050022
2.中国科学院大学 北京 100049
3.青海师范大学生命与地理科学学院 西宁 810000
基金项目: 国家重点研发计划课题2016YFC0401403
国家自然科学基金面上项目31870422
河北省自然科学基金项目D2016503001
中国科学院青年创新促进会项目2017138

详细信息

作者简介:姜寒冰, 主要从事作物水分利用效率和农田水平衡过程研究。E-mail:jianghanbing16@163.com

通讯作者:沈彦俊, 主要从事生态水文过程的研究。E-mail:yjshen@sjziam.ac.cn

中图分类号:S181

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收稿日期:2018-05-25
录用日期:2018-09-02
刊出日期:2019-01-01

A review of progress in research and scaling-up methods of crop water use efficiency

JIANG Hanbing^{1, 2,},
ZHANG Yucui¹,
REN Xiaodong^{1, 3},
YAO Jiawei^{1, 2},
SHEN Yanjun^1,,
1. Key Laboratory of Agricultural Water Resources, Chinese Academy of Sciences/Hebei Laboratory of Water-Saving Agriculture/Center for Agricultural Resources Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Shi-jiazhuang 050022, China
2. University of Chinese Academy of Sciences, Beijing 100049, China
3. College of Biological and Geo-graphical Sciences, Qinghai Normal University, Xining 810000, China
Funds: the National Key Research and Development Plan of China2016YFC0401403
the National Natural Science Foundation of China31870422
the Natural Science Foundation of Hebei ProvinceD2016503001
the Youth Innovation Promotion Association of Chinese Academy of Sciences2017138

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Corresponding author:SHEN Yanjun, E-mail:yjshen@sjziam.ac.cn

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摘要
摘要:提高作物水分利用效率（WUE）是缓解农业生产水资源匮乏压力的有效途径，而水分利用效率尺度传递是各尺度WUE相互表征、验证并应用于实际生产的基础。本文概述了作物叶片、植株、群体尺度WUE的主要观测技术，包括叶片气体交换测定、碳同位素判别、桶栽称重、涡度相关观测等，其中碳同位素判别法为研究作物水分利用的长期累积效应提供了新的思路，且适用于多个尺度；总结了各尺度WUE的影响因子及作物耗水的生理机制，阐明各尺度WUE均受气孔导度调控。讨论了叶片到植株、叶片/植株到群体的尺度传递的可行性，集中分析了尺度传递的主要限制因素，指出叶片到植株的传递研究难点集中于叶片分布和光分布的不确定性、植物夜间呼吸和蒸腾以及植物适应环境的生理调节机制等过程；而叶片/植株到群体的传递研究主要受冠层形态学差异、冠层阻力、土壤蒸发及植物同化物分配机制等限制。最后总结了尺度传递方法的现有研究成果。目前作物WUE尺度传递主要依靠模型的完善和观测手段的提高，叶片到单株的尺度传递需关注日间与夜间耗水的分离及作物各部分的光合特性；叶片/单株到群体的传递可先明确蒸散结构，了解耗水特征，再以气孔导度和冠层导度的关系为切入点，利用模型探究传递机制。
关键词:水分利用效率/
尺度传递/
碳同位素/
气孔导度
Abstract:Increasing crop water use efficiency (WUE) is an effective way of alleviating agricultural water scarcity. The scaling up of water use efficiency is the basis for mutual representation, verification and application of achievements at various scales. This paper summarized the main observation technologies of leaf-scale, plant-scale and plantation-scale WUE. At present, the widely used methods include leaf gas exchange measurement, carbon isotopic discrimination, pot weighing method, eddy covariance system, etc. Carbon isotope discrimination provides a new idea for the study of long-term cumulative effects of crop water use conditions which is also available at every scale. We reviewed the impacting factors and the related physiological mechanisms of crop water use at multi-scale WUE. Crop WUE at each scale was regulated by stomatal conductance and crops usually regulated stomatal aperture to response to temperature, humidity, CO₂ and other interactive environmental factors. Stomatal optimization theory essentially sought optimal state of stomata under complex environmental conditions to coordinate the process of photosynthesis and transpiration of crops. Instantaneous WUE at leaf scale cannot directly represent water use status at larger spatial and temporal scales. Thus we also discussed the feasibility of scaling up WUE from leaf to plant to plantation scales and analyzed the main limiting factors at each scale transfer. We pointed out the difficulties in transfer from leaf to plant in terms of WUE. It mainly focused on three points-uncertainty in leaf and light distributions, plant nighttime respiration and transpiration, and plant physiological adjustment mechanisms. Research on leaf to plant to plantation scale transfer was mainly influenced by canopy internal resistance, boundary layer resistance, soil evaporation, night transpiration of crops, crop water use and assimilates partitioning mechanism. Finally, existing research achievements on scale transfer were summarized. At present, WUE scale transfer depended mainly on improvement of models and observation methods. The transfer from leaf to plant focused on separation of water use during day and night and photosynthetic characteristics of each part of the crop. For transfer from leaf to plant to plantation scale, studies explored efficient ways. First, studies understood the structure of evapotranspiration and confirmed the characteristics of water use. Second, studies used the relationship between stomatal and canopy conductance as breakthrough point via models to explore transfer mechanisms. Actually, several models had already been established and applied in this respect.
Key words:Water use efficiency/
Scaling up/
Carbon isotopic discrimination/
Stomatal conductance

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表1不同尺度水分利用效率(WUE)的定义与观测手段
Table1.Definition and measurement methods of water use efficiency (WUE) under different scales

尺度 Scale	WUE	计算公式 Formula	观测手段 Measurement method	应用意义 Application
叶片 Single leaf	WUET	A/T	气体交换法, 稳定碳同位素技术[6] Gas exchange measurement, stable carbon isotope technology[6]	作物生理状态表征[7-8] Plant ecophysiology characterization[7-8]
	WUEi	A/gs
植株 Single plant	WUEp	TB/T	称重法、稳定碳同位素技术[9] Pot weighting method, stable carbon isotope technology[9]	作物育种优良单株选择[10] Superior plant selection in crop breeding[10]
群体 Population	WUEc	GPP/ET; NPP/ET; NEE/ET	涡度相关技术、遥感监测、农田水平衡[11-13] Eddy Covariance Technology, remote sensing monitor, field water balance[11-13]	生态系统综合评价[14] Ecology system assessment[14]
	WUEf	TB/ET; Yield/ET
WUET:瞬时水分利用效率; WUEi:内部水分利用效率; WUEp:植株尺度水分利用效率; WUEc:冠层水平水分利用效率; WUEf:农田尺度水分利用效率; A:净CO2吸收速率; T:蒸腾速率; gs:气孔导度; TB:干物质量; GPP:初级生产力; NPP:净初级生产力; NEE:净生态系统碳交换量; ET:蒸散耗水量。Yield:产量。WUET: instantaneous WUE; WUEi: internal WUE; WUEp: WUE at plant scale; WUEc: WUE at canopy level; WUEf: WUE at field scale. A: net CO2 absorption rate; T: transpiration rate; gs: stomatal conductance; TB: dry biomass weight; GPP: gross primary productivity; NPP: net primary productivity; NEE: net ecosystem carbon exchange; ET: evapotranspiration.

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表2不同尺度作物水分利用效率及尺度传递研究成果
Table2.Achievements of water use efficiency (WUE) of crops and scale-up methods under different scales

研究尺度 Scale	关键因素 Vital factors	主要成果 Primary achievement	文献和年份 Reference and year
叶片(已应用于多个尺度) Single leaf (applied to multiple scales)	多环境因子 Multiple environmental factors	水碳耦合模型(已发展出BWB模型, BBL模型等) Water-carbon coupling model (BWB model, BBL model, et al)	[40] 1988
叶片 Single leaf	边界层导度 Boundary layer conductance	光合-蒸腾-气孔导度耦合模型 Integrated photosynthesis-transpiration-stomatal conductance model	[41] 1998
叶片 Single leaf	叶肉细胞导度、气孔行为 Mesophyll conductance; stomatal behavior	SMPT-SB模型 SMPT-SB model	[42] 2003
叶片-冠层 From single leaf to canopy	光合有效辐射、饱和水汽压差 Photosynthetically active radiation (PAR), water vapor deficit (VPD)	构建气孔导度到冠层导度的尺度拓展估算模型 A model scaling up from leaf stomatal conductance to canopy conductance	[43] 2011
叶片、单株 Single leaf, single plant	水分条件 Water condition	适度水分亏缺下, 叶片气孔导度可作为单株WUE的指示因子 Stomatal conductance of leaves can be used as an indicator of the whole-plant WUE under moderate water deficit	[25] 2012
叶片、单株 Single leaf, single plant	暗处理 Dark treatment	暗处理下的叶片气孔导度比枯萎叶片的最小导度更能指示单株 WUE Dark-adapted leaf stomatal conductance, but not minimum leaf conductance of wilted leaves, predicts whole-plant WUE	[44] 2013
叶片-冠层 From single leaf to canopy	光合有效辐射(阴生叶、阳生叶) PAR (sun-leaf; shading-leaf)	双源双叶模型 Dual-source dual-leaf model	[45] 2014
叶片、单株 Single leaf, single plant	饱和水汽压差、叶片位置 VPD; leaf position	叶片向单株拓展的主要限制因素 Main limitations of scaling up from single leaf to plant	[4] 2015
叶片、单株、冠层 Single leaf, single, plant canopy	多环境因子 Multiple environmental factors	各尺度观测结果的不确定性 Uncertainty of measurements at multi-scales	[46] 2017
叶片、单株 Single leaf, single plant	叶片水势 Leaf water potential	利用植物结构功能模型模拟优化不同冠层形状的 WUE WUE simulation of different canopy type with Functional-Structural Plant Models	[47] 2016

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