植物水分调节对策: 等水与非等水行为

doi:10.17521/cjpe.2016.0366

摘要/Abstract

摘要：

水分是植物存活、生长和分布过程中的必需资源, 阐明植物对干旱的应对和调节机制, 是植物生理生态学和全球变化生态学的重要研究命题。植物对不同气候与土壤水分条件的长期适应会形成由一整套相关联的性状组成的水分调节策略, 其中等水和非等水调节行为是两种典型的水分调节对策。区分并阐明植物的水分调节对策及其机制, 不但在干旱地区植物育种、植被修复等实践中有广泛的应用前景, 而且可为构建更精确的植被动态模型和预测气候变化情景下植被分布提供科学基础。该文首先阐述了等水和非等水调节行为的定义及3种定量分类方法: (1)基于气孔导度与叶水势的关系; (2)基于气孔导度与水汽压亏缺的关系; (3)基于黎明前叶水势与中午叶水势的关系。之后, 从水力和碳经济性状两个方面比较分析了两种水分调节对策植物的种间差异。综合分析植物水分调节机制发现, 水力信号与化学信号的相互作用是植物水分调节行为的主控因素。最后提出3个亟待开展研究的问题: (1)针对不同地区开展植物水分关系相关性状的测定, 寻求可靠且普适的植物水分调节对策分类方法。(2)探索植物水分调节对策与水力、形态、结构、功能等性状之间的关联性, 为改进植被动态模型提供可靠的参数。(3)加深理解不同时空尺度上植物水分调节过程, 揭示植物对环境胁迫(尤其是干旱)的响应和适应机制。

关键词: 干旱胁迫, 木质部栓塞, 气候变化, 气孔调节, 水力失衡, 植物性状

Abstract:

Water is a vital resource for plant survival, growth and distribution, and it is of significance to explore mechanisms of plant water-relations regulation and responses to drought in ecophysiology and global change ecology. Plants adapt to different climates and soil water regimes and develop divergent water-regulation strategies involving a suite of related traits, of which two typical types are isohydric and anisohydric behaviors. It is critical to distinguish water-regulation strategies of plants and reveal the underlying mechanisms for plant breeding and vegetation restoration especially in xeric regions; and it is also important for developing more accurate vegetation dynamic models and predicting vegetation distribution under climate change scenarios. In this review, we first recalled the definitions of isohydric and anisohydric regulations and three quantitative classification methods that were established based on the relationships (1) between stomatal conductance and leaf water potential, (2) between stomatal conductance and vapor pressure deficit, (3) between predawn and midday leaf water potentials. We then compared the two water-regulation strategies in terms of hydraulics and carbon-economics traits. We synthesized the mechanisms of plant water-regulation and found that the interaction between hydraulic and chemical signals was the dominant factor controlling plant water-regulation behavior. Last, we proposed three promising aspects in this field: (1) to explore reliable and universal methods for classifying plant water-regulation strategies based on extensive investigation of the traits related with plant water-relations in various regions; (2) to explore relationships between plant water-regulation strategies and traits of hydraulics, morphology, structure, and function in order to provide reliable parameters for improving vegetation dynamic models; and (3) to deeply understand the processes of plant water-regulation at different spatial and temporal scales, and reveal mechanisms of plants’ responses and adaption to environmental stresses (especially drought).

Key words: drought stress, xylem embolism, climate change, stomatal regulation, hydraulic failure, plant trait

罗丹丹, 王传宽, 金鹰. 植物水分调节对策: 等水与非等水行为. 植物生态学报, 2017, 41(9): 1020-1032. DOI: 10.17521/cjpe.2016.0366

Dan-Dan LUO, Chuan-Kuan WANG, Ying JIN. Plant water-regulation strategies: Isohydric versus anisohydric behavior. Chinese Journal of Plant Ecology, 2017, 41(9): 1020-1032. DOI: 10.17521/cjpe.2016.0366

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https://www.plant-ecology.com/CN/Y2017/V41/I9/1020

图/表 2

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93	Appendix I Terms and their acronyms or symbol

性状 Trait		等水调节 Isohydric regulation	非等水调节 Anisohydric regulation	是否存在争议 Challenged or not
水力性状 Hydraulics	生长策略 Growth strategy	保守型 Conservative behaviour	冒险型 Risk-taking behaviour	N
	最小叶水势 Minimum leaf water potential	相对恒定(高) Constant (High)	低 Low	N
	气孔导度 Stomatal conductance	低 Low	相对恒定(高) Constant (High)	Y (Quero et al., 2011)
	导水率 Hydraulic conductance	低 Low	高 High	N
	耐旱性 Drought tolerance	弱 Weak	强 Strong	Y (Quero et al., 2011)
	木质部脆弱性 Xylem vulnerability	小 Small	大 Large	N
	水力安全阈值 Safety margin	大 Large	小 Small	N
	栓塞恢复力 Embolism recovery ability	弱 Weak	强 Strong	Y (McCulloh & Meinzer., 2015)
	纹孔膜 Pit membrane	厚; 总面积小 Thick; Smaller total area	薄; 总面积大 Thin; Larger total area	-
碳经济性状 Carbon economics	光合速率 Photosynthetic rate	小 Small	大 Large	Y (Quero et al., 2011)
	呼吸速率 Respiratory rate	小 Small	大 Large	N
	内在水分利用效率 Intrinsic water use efficiency	高 High	低 Low	Y (Lovisolo et al., 2010)
	非结构性碳水化合物 Nonstructural carbohydrate	低 Low	高 High	Y (Woodruff et al., 2015)
	比叶质量 Leaf mass per area	大 Large	小 Small	-
	叶寿命 Leaf lifespan	长 Long	短 Short	-

性状 Trait		等水调节 Isohydric regulation	非等水调节 Anisohydric regulation	是否存在争议 Challenged or not
水力性状 Hydraulics	生长策略 Growth strategy	保守型 Conservative behaviour	冒险型 Risk-taking behaviour	N
	最小叶水势 Minimum leaf water potential	相对恒定(高) Constant (High)	低 Low	N
	气孔导度 Stomatal conductance	低 Low	相对恒定(高) Constant (High)	Y (Quero et al., 2011)
	导水率 Hydraulic conductance	低 Low	高 High	N
	耐旱性 Drought tolerance	弱 Weak	强 Strong	Y (Quero et al., 2011)
	木质部脆弱性 Xylem vulnerability	小 Small	大 Large	N
	水力安全阈值 Safety margin	大 Large	小 Small	N
	栓塞恢复力 Embolism recovery ability	弱 Weak	强 Strong	Y (McCulloh & Meinzer., 2015)
	纹孔膜 Pit membrane	厚; 总面积小 Thick; Smaller total area	薄; 总面积大 Thin; Larger total area	-
碳经济性状 Carbon economics	光合速率 Photosynthetic rate	小 Small	大 Large	Y (Quero et al., 2011)
	呼吸速率 Respiratory rate	小 Small	大 Large	N
	内在水分利用效率 Intrinsic water use efficiency	高 High	低 Low	Y (Lovisolo et al., 2010)
	非结构性碳水化合物 Nonstructural carbohydrate	低 Low	高 High	Y (Woodruff et al., 2015)
	比叶质量 Leaf mass per area	大 Large	小 Small	-
	叶寿命 Leaf lifespan	长 Long	短 Short	-

缩写或符号 Acronym or symbol	术语 Term
AAO	脱落醛氧化酶 Abscisic aldehyde oxidase
ABA	脱落酸 Abscisic acid
A_N	净CO₂同化量 Net CO₂ assimilation
AQPs	水通道蛋白 Aquaporins
G_s	气孔导度 Stomatal conductance
T_r	蒸腾速率 Transpiration rate
K	水力导度 Hydraulic conductance
K_leaf	叶水力导度 Leaf hydraulic conductance
K_plant	植株导水率 Whole-plant hydraulic conductivity
LMA	比叶质量 Leaf dry mass per area
MCSU	钼辅因子硫化酶 Molybdate cofactor sulfurase
NCED	9-顺式-环氧类胡萝卜素加双氧酶蛋白 9-cis-epoxycarotenoid dioxygenase
NSC	非结构性碳水化合物 Nonstructural carbohydrate
P₅₀	木质部失去50%导水率所对应的水势 The water potential inducing 50% loss of hydraulic conductivity
P₈₈	木质部失去88%导水率所对应的水势 The water potential inducing 88% loss of hydraulic conductivity
Pe	栓塞临界值 Embolism threshold
TIP	液泡膜内在蛋白 Tonoplast-intrinsic protein
VPD	水汽压亏缺 Vapor pressure deficit
WUE	水分利用效率 Water use efficiency
WUE_i	内在水分利用效率 Intrinsic water use efficiency
ZEP	玉米黄质环氧酶 Zeaxanthin epoxidase
Ψ_L	叶水势 Leaf water potential
Ψ_MD	中午叶水势 Midday leaf water potential
Ψ_PD	黎明前叶水势 Predawn leaf water potential
Ψ_S	土壤水势 Soil water potential

缩写或符号 Acronym or symbol	术语 Term
AAO	脱落醛氧化酶 Abscisic aldehyde oxidase
ABA	脱落酸 Abscisic acid
A_N	净CO₂同化量 Net CO₂ assimilation
AQPs	水通道蛋白 Aquaporins
G_s	气孔导度 Stomatal conductance
T_r	蒸腾速率 Transpiration rate
K	水力导度 Hydraulic conductance
K_leaf	叶水力导度 Leaf hydraulic conductance
K_plant	植株导水率 Whole-plant hydraulic conductivity
LMA	比叶质量 Leaf dry mass per area
MCSU	钼辅因子硫化酶 Molybdate cofactor sulfurase
NCED	9-顺式-环氧类胡萝卜素加双氧酶蛋白 9-cis-epoxycarotenoid dioxygenase
NSC	非结构性碳水化合物 Nonstructural carbohydrate
P₅₀	木质部失去50%导水率所对应的水势 The water potential inducing 50% loss of hydraulic conductivity
P₈₈	木质部失去88%导水率所对应的水势 The water potential inducing 88% loss of hydraulic conductivity
Pe	栓塞临界值 Embolism threshold
TIP	液泡膜内在蛋白 Tonoplast-intrinsic protein
VPD	水汽压亏缺 Vapor pressure deficit
WUE	水分利用效率 Water use efficiency
WUE_i	内在水分利用效率 Intrinsic water use efficiency
ZEP	玉米黄质环氧酶 Zeaxanthin epoxidase
Ψ_L	叶水势 Leaf water potential
Ψ_MD	中午叶水势 Midday leaf water potential
Ψ_PD	黎明前叶水势 Predawn leaf water potential
Ψ_S	土壤水势 Soil water potential