Interactive effects of ozone and drought stress on plants: A review

doi:10.17521/cjpe.2016.0195

Abstract

Abstract:

Ground-level ozone (O₃) and drought are two key factors limiting plant growth. O₃ can enter into the plant tissue through the stomata, then causing the formation of reactive oxygen species (ROS) which inspires programmed cell death. Drought usually induces the accumulation of ROS due to damage to antioxidant systems of plants. The effects of two kinds of stress on plants are similar due to the accumulation of ROS, resulting in reduced photosynthesis rate and physiological metabolism, eventually decreased plant growth and biomass. Nevertheless, O₃and drought interacts synergistically to accumulate detrimental effects or antagonistically to reduce harmful effects. Actually, it is complex interactive process between O₃ and drought. On the one hand, O₃ triggers stomatal sluggishness or even dysfunction, which exacerbates water transpiration of leaves, water loss from plants and further O₃ phytotoxicity. On the other hand, drought induces stomatal closure, and thus protecting plants against the O₃ influx and evaporation of water. However, prolonged drought could limit the uptake of CO₂ and thus result in reduced plant growth. The response of plants to both O₃ and drought not only depends on the occurring sequence and duration of any factor but also rely on the difference in physiological metabolism of the plant itself. The interactive effects of O₃ and drought on stomatal characteristics, photosynthetic carbon mechanism, antioxidant response and growth development are reviewed in this paper and the aspects to be further studied are also suggested.

Key words: O₃, drought, interaction, plant, response

Feng GAO, Pin LI, Zhao-Zhong FENG. Interactive effects of ozone and drought stress on plants: A review[J]. Chin J Plan Ecolo, 2017, 41(2): 252-268.

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植物类型 Plant type	植物种类 Plants species	树龄(年) Tree-age (year)	实验方法Experimental method	O₃处理 O₃ treatment (nmol·mol^-1)	干旱处理 Drought treatment	研究内容 Research content	参考文献 Reference
草本Herbaceous	冬小麦 Triticum aestivum	-	温室 Greenhouse	80	SWC 60%, 45%, 35%	叶片O₃吸收与产量 Leaf ozone uptake and grain yield	Khan & Soja, 2003
	冬小麦 Triticum aestivum	-	OTC	125	SWC 35%-40%	生理生长 Physiology and biochemistry	Xu et al., 2007
	冬小麦 Triticum aestivum, 圆锥小麦 Triticum turgidum ssp. durum	-	OTC	83	SWC 42%	生理生化、产量 Physiology and biochemistry, grain yield	Biswas & Jiang, 2011
	春小麦 Triticum aestivum, Triticum durum	-	OTC	AA + 50	40%对照SWC 40% SWC of control	抗氧化系统 Antioxidant defense systems	Herbinger et al., 2002
	大豆 Glycine max	-	OTC	0.70 × AA, 60	-0.8MPa黎明前叶片水势 - 0.8 MPa Pre-dawn leaf water potential	蒸散 Evapotranspiration	Rana et al., 2012
	菜豆 Pharsalus vulgaris	-	GC	48, 87, 150	SWC 6%-15%	气孔 Stomata	Hoshika et al., 2013
	番茄 Lycopersicon esculentum	-	大田 Field	AA, AA + 80	18%-33%田间持水量 18%-33% Field capacity	果实自由基含量 Radical contents in fruit	Pirker et al., 2003
	Leontodon hispidus	-	GC	70	50%田间持水量 50% Field capacity	脱落酸(ABA)作用机制 Mechanisms of action of abscisic acid	Wilkinson & Davies, 2009
	Medicago runcatula	-	GC	70	以测定量灌溉 Adding measured amounts of water	生理生化、分子 Physiological, biochemical and molecular response	Iyer et al., 2013
	烟草 Nicotiana tabacum	-	GC	200	适当灌溉 With holding water from the plants	分子(DHAR表达) Molecular (dehydroascorbate reductase overexpressing)	Eltayeb et al., 2006
常绿、落叶乔木 Evergreen and deciduous trees	欧洲云杉 Picea abies, Fagus sylvatica	50; 60	开放式熏气系统 A free-air fumigation system	2 × AA	NE (2年不同气候) NE (Climatic variation in two years)	地下影响(土壤呼吸、根系生物量) Belowground effects (soil respiration, fine-root biomass)	Nikolova et al., 2010
	欧洲云杉 Picea abies, Fagus sylvatica	-	NE	AA	NE (两种树分别12个和4个不同地点) NE (12 and 4 different site conditions were used, respectively)	规模性生长关系 Size-growth relationship	Pretzsch & Dieler, 2011
	阿巴拉契亚山脉南部森林 Southern Appalachian forest	成熟林 Mature forest trees	NE	AA	NE (3个不同地点在2-3年的不同气候) (Three sites over 2-3 years)	生长和液流速率 Growth and sap flow velocity	McLaughlin et al., 2007
常绿乔木 Evergreen trees	欧洲云杉 Picea abies	4	OTC	37	胁迫期停止灌溉 The irrigation system was turned off during the dry periods	叶肉细胞显微结构 Microscopic structures of mesophyll cells	Kivimäenpää et al., 2001
	欧洲云杉 Picea abies	2-5	OTC	26-30, 25-29, 29-33	以土壤水势确定灌溉水量 Watered by the soil water potential	叶绿素、养分和生长 Chlorophyll, mineral nutrients and growth	Wallin et al., 2002
	欧洲云杉 Picea abies	2-5	OTC	1.5 × AA	以土壤水势确定灌溉水量 Watered by the soil water potential	叶片细胞结构 Cell structural changes in the needles	Kivimäenpää et al., 2003
	Abies veitchii	6	GC	50	土壤水势6.19-30.96 kPa; 30.96-97.98 kPa Soil water potential 6.19-30.96 kPa; 30.96-97.98 kPa	生长 Growth	Feng & Shimizu, 2005
	Abies concolor	2; 53	OTC, NE	52, 30-35	0, 10%, 25%, 50%的年降水量减少 0, 10%, 25%, 50% Reductions in total annual precipitation	模型模拟胁迫下生长 Simulation growth by model under stress	Retzlaff et al., 2000
	Pinus halepensis	2	OTC	69	50%对照灌溉水量 Half the water supplied to control	叶片抗氧化酶类 Antioxidative enzymes in needles	Alonso et al., 2001
	Pinus halepensis	4	GC	300	以叶片水势和气体交换临界值确定灌水量 Characterized by very low leaf water potential and gas exchange values	叶片生理 Physiological response of needles	Manes et al., 2001
植物类型 Plant type	植物种类 Plants species	树龄(年) Tree-age (year)	实验方法Experimental method	O₃处理 O₃ treatment (nmol·mol^-1)	干旱处理 Drought treatment	研究内容 Research content	参考文献 Reference
常绿乔木 Evergreen trees	Pinus halepensis	3	GC	200	100%-50%缓慢水分散失(中度); 以黎明前叶片水势确定灌溉水量(重度) From 100% to 50% of the water loss by evapotranspiration (mild); watered by pre-dawn needle water potential (intensity)	叶片酶(PEPC) Phosphoenolpyruvatecarboxylase in needles	Fontaine et al., 2003
	Pinus halepensis	2	OTC	AA + 40	50%对照灌溉水量 Half the water supplied to control	生理生化补偿过程 Compensation processes in physiology and biochemistry	Inclan et al., 2005
	Quercus ilex	3	气候室 Climatic chambers	250	20 mL水每间隔一周灌溉 20 mL of water per plant every week	生理响应 Physiological responses	Vitale et al., 2008
	Quercus ilex ssp. ilex, 长角豆 Ceratonia siliqua	1	OTC	37; 57	50%对照灌溉次数 Half the watered times to control	生理生化 Physiology and biochemistry	Ribas et al., 2005
	Quercus ilex ssp. ilex, Q. ilex ssp. ballota	1-2	OTC	31; 49	50%对照灌溉水量 Half the water supplied to control	气体交换、生长和生物量 Gas exchange, growth and biomass	Alonso et al., 2014
	西黄松 Pinus ponderosa	6-8	NE	40-70	NE	O₃和碳吸收 O₃and carbon uptake	Panek & Goldstein, 2001
	西黄松 Pinus ponderosa	40	NE	58-60, 64-67, 80	黎明前木质部水势< -1.2 MPa Xylem potentials <-1.2 MPa at predawn	碳获取 Carbon acquisition	Grulke et al., 2002
	西黄松 Pinus ponderosa	5-7	NE	53	NE (2年不同气候) NE (Climatic variation in two years)	O₃暴露度量参数 O₃ exposure metrics parameters	Panek et al., 2002
	西黄松 Pinus ponderosa	7	NE	AA	NE (4个不同地点) NE (Four sites)	吸收模型参数 Parameters for O₃ uptake modeling	Panek, 2004
落叶乔木Deciduous trees	海南蒲桃 Syzygium hainanense, 糖胶树 Alstonia scholaris	当年 Current year	OTC	75	40%-50%土壤相对含水量 40%-50% Relative soil water content	光合生理 Photosynthetic physiology	Hao et al., 2014
	长芒杜英 Elaeocarpus apiculatus, 壳菜果Mytilaria laosensis,黧蒴锥 Castanopsis fissa 醉香含笑 Michelia macclurei, 樟 Cinnamomum camphora, 红花荷Rhodoleia championii, 壳菜果 Mytilaria laosensis	当年 Current year 1	OTC OTC	150 20	50%对照灌溉次数 Half the watered times to control 两天一次灌溉 Watered once in two days	荧光生理 Chlorophyll a fluorescence 根、茎、叶和总生物量及根茎比 Root biomass, stem biomass, leaf biomass, total biomass and root/shoot ratio	Li et al., 2015 Ye et al., 2014
	Fagus crenata	3	GC	60	70%对照灌溉水量 70% Water supplied to control	叶片物候和芽抗寒性 Leaf phenological characteristics and bud frost hardiness	Yonekura et al., 2004
	Fagus crenata	3	GC	60	70%对照灌溉水量 70% Water supplied to control	叶片抗氧化系统 Leaf antioxidative systems	Watanabe et al., 2005
	Fagus sylvatica	60	NE	2 × AA	NE (3年不同气候) NE (Climatic variation in three years)	生理生化 Physiology and biochemistry	Löw et al., 2006
	Fagus sylvatica	60	NE	2 × AA	NE (2年不同气候) NE (Climatic variation in two years)	细根抗氧化物 Antioxidants in fine roots	Haberer et al., 2008
	Fagus sylvatica	-	NE	AA	NE (9个不同地点) NE (Nine sites)	日生长 Daily growth	Kuehn et al., 2015
	Fagus sylvatica, 夏栎 Quercus robur, 辽杨×中东杨 Populus maximowiczii × P. berolinensis	1	OTC	0.95 × AA	50%田间持水量 50% Field capacity	生长和生理响应 Growth and physiological responses	Pollastrini et al., 2010
	岳桦 Betula ermanii	2	GC	50	土壤水势 6.19-30.96 kPa, 30.96-97.98 kPa Soil water potential 6.19-30.96 kPa, 30.96-97.98 kPa	生长、生理生化 Growth、physiology and biochemistry	Shimizu & Feng, 2007
	元宝槭 Acer truncatum	1	OTC	102-147	40%-50%田间持水量 40%-50% Field capacity	气孔响应 Stomatal response	Wen et al., 2014
	元宝槭 Acer truncatum	1	OTC	102-147	40%-50%田间持水量 40%-50% Field capacity	生长和生理 Growth and physiology	Li et al., 2015
植物类型 Plant type	植物种类 Plants species	树龄(年) Tree-age (year)	实验方法Experimental method	O₃处理 O₃ treatment (nmol·mol^-1)	干旱处理 Drought treatment	研究内容 Research content	参考文献 Reference
落叶乔木Deciduous trees	辽杨×中东杨 Populus maximowiczii × P. berolinensis	当年 Current year	OTC	0.95 × AA	150 mL水每天灌溉 150 mL of water a day	不同冠层叶绿素荧光 Chlorophyll a fluorescence along a crown	Desotgiu et al., 2012
	辽杨×中东杨 Populus maximowiczii × P. berolinensis	当年 Current year	OTC	0.95 × AA	150 mL水每天灌溉 150 mL water a day	光合、生长和同位素 Photosynthesis、growth and stable isotope	Pollastrini et al., 2013
	欧洲山杨×银白杨 Populus tremula × P. alba	当年 Current year	GC	CF + 120	SWC 35%	生理、蛋白质组 Physiological and proteomic	Bohler et al., 2013
	辽杨×中东杨 Populus maximowiczii × P. berolinensis	当年 Current year	OTC	44-53	60%, 20%对照灌溉水量 60%, 20% Water respect to control	生理生化、生长和同位素 Physiological and biochemistry、biomass and isotope	Pollastrini et al., 2014

植物类型 Plant type	植物种类 Plants species	树龄(年) Tree-age (year)	实验方法Experimental method	O₃处理 O₃ treatment (nmol·mol^-1)	干旱处理 Drought treatment	研究内容 Research content	参考文献 Reference
草本Herbaceous	冬小麦 Triticum aestivum	-	温室 Greenhouse	80	SWC 60%, 45%, 35%	叶片O₃吸收与产量 Leaf ozone uptake and grain yield	Khan & Soja, 2003
	冬小麦 Triticum aestivum	-	OTC	125	SWC 35%-40%	生理生长 Physiology and biochemistry	Xu et al., 2007
	冬小麦 Triticum aestivum, 圆锥小麦 Triticum turgidum ssp. durum	-	OTC	83	SWC 42%	生理生化、产量 Physiology and biochemistry, grain yield	Biswas & Jiang, 2011
	春小麦 Triticum aestivum, Triticum durum	-	OTC	AA + 50	40%对照SWC 40% SWC of control	抗氧化系统 Antioxidant defense systems	Herbinger et al., 2002
	大豆 Glycine max	-	OTC	0.70 × AA, 60	-0.8MPa黎明前叶片水势 - 0.8 MPa Pre-dawn leaf water potential	蒸散 Evapotranspiration	Rana et al., 2012
	菜豆 Pharsalus vulgaris	-	GC	48, 87, 150	SWC 6%-15%	气孔 Stomata	Hoshika et al., 2013
	番茄 Lycopersicon esculentum	-	大田 Field	AA, AA + 80	18%-33%田间持水量 18%-33% Field capacity	果实自由基含量 Radical contents in fruit	Pirker et al., 2003
	Leontodon hispidus	-	GC	70	50%田间持水量 50% Field capacity	脱落酸(ABA)作用机制 Mechanisms of action of abscisic acid	Wilkinson & Davies, 2009
	Medicago runcatula	-	GC	70	以测定量灌溉 Adding measured amounts of water	生理生化、分子 Physiological, biochemical and molecular response	Iyer et al., 2013
	烟草 Nicotiana tabacum	-	GC	200	适当灌溉 With holding water from the plants	分子(DHAR表达) Molecular (dehydroascorbate reductase overexpressing)	Eltayeb et al., 2006
常绿、落叶乔木 Evergreen and deciduous trees	欧洲云杉 Picea abies, Fagus sylvatica	50; 60	开放式熏气系统 A free-air fumigation system	2 × AA	NE (2年不同气候) NE (Climatic variation in two years)	地下影响(土壤呼吸、根系生物量) Belowground effects (soil respiration, fine-root biomass)	Nikolova et al., 2010
	欧洲云杉 Picea abies, Fagus sylvatica	-	NE	AA	NE (两种树分别12个和4个不同地点) NE (12 and 4 different site conditions were used, respectively)	规模性生长关系 Size-growth relationship	Pretzsch & Dieler, 2011
	阿巴拉契亚山脉南部森林 Southern Appalachian forest	成熟林 Mature forest trees	NE	AA	NE (3个不同地点在2-3年的不同气候) (Three sites over 2-3 years)	生长和液流速率 Growth and sap flow velocity	McLaughlin et al., 2007
常绿乔木 Evergreen trees	欧洲云杉 Picea abies	4	OTC	37	胁迫期停止灌溉 The irrigation system was turned off during the dry periods	叶肉细胞显微结构 Microscopic structures of mesophyll cells	Kivimäenpää et al., 2001
	欧洲云杉 Picea abies	2-5	OTC	26-30, 25-29, 29-33	以土壤水势确定灌溉水量 Watered by the soil water potential	叶绿素、养分和生长 Chlorophyll, mineral nutrients and growth	Wallin et al., 2002
	欧洲云杉 Picea abies	2-5	OTC	1.5 × AA	以土壤水势确定灌溉水量 Watered by the soil water potential	叶片细胞结构 Cell structural changes in the needles	Kivimäenpää et al., 2003
	Abies veitchii	6	GC	50	土壤水势6.19-30.96 kPa; 30.96-97.98 kPa Soil water potential 6.19-30.96 kPa; 30.96-97.98 kPa	生长 Growth	Feng & Shimizu, 2005
	Abies concolor	2; 53	OTC, NE	52, 30-35	0, 10%, 25%, 50%的年降水量减少 0, 10%, 25%, 50% Reductions in total annual precipitation	模型模拟胁迫下生长 Simulation growth by model under stress	Retzlaff et al., 2000
	Pinus halepensis	2	OTC	69	50%对照灌溉水量 Half the water supplied to control	叶片抗氧化酶类 Antioxidative enzymes in needles	Alonso et al., 2001
	Pinus halepensis	4	GC	300	以叶片水势和气体交换临界值确定灌水量 Characterized by very low leaf water potential and gas exchange values	叶片生理 Physiological response of needles	Manes et al., 2001
植物类型 Plant type	植物种类 Plants species	树龄(年) Tree-age (year)	实验方法Experimental method	O₃处理 O₃ treatment (nmol·mol^-1)	干旱处理 Drought treatment	研究内容 Research content	参考文献 Reference
常绿乔木 Evergreen trees	Pinus halepensis	3	GC	200	100%-50%缓慢水分散失(中度); 以黎明前叶片水势确定灌溉水量(重度) From 100% to 50% of the water loss by evapotranspiration (mild); watered by pre-dawn needle water potential (intensity)	叶片酶(PEPC) Phosphoenolpyruvatecarboxylase in needles	Fontaine et al., 2003
	Pinus halepensis	2	OTC	AA + 40	50%对照灌溉水量 Half the water supplied to control	生理生化补偿过程 Compensation processes in physiology and biochemistry	Inclan et al., 2005
	Quercus ilex	3	气候室 Climatic chambers	250	20 mL水每间隔一周灌溉 20 mL of water per plant every week	生理响应 Physiological responses	Vitale et al., 2008
	Quercus ilex ssp. ilex, 长角豆 Ceratonia siliqua	1	OTC	37; 57	50%对照灌溉次数 Half the watered times to control	生理生化 Physiology and biochemistry	Ribas et al., 2005
	Quercus ilex ssp. ilex, Q. ilex ssp. ballota	1-2	OTC	31; 49	50%对照灌溉水量 Half the water supplied to control	气体交换、生长和生物量 Gas exchange, growth and biomass	Alonso et al., 2014
	西黄松 Pinus ponderosa	6-8	NE	40-70	NE	O₃和碳吸收 O₃and carbon uptake	Panek & Goldstein, 2001
	西黄松 Pinus ponderosa	40	NE	58-60, 64-67, 80	黎明前木质部水势< -1.2 MPa Xylem potentials <-1.2 MPa at predawn	碳获取 Carbon acquisition	Grulke et al., 2002
	西黄松 Pinus ponderosa	5-7	NE	53	NE (2年不同气候) NE (Climatic variation in two years)	O₃暴露度量参数 O₃ exposure metrics parameters	Panek et al., 2002
	西黄松 Pinus ponderosa	7	NE	AA	NE (4个不同地点) NE (Four sites)	吸收模型参数 Parameters for O₃ uptake modeling	Panek, 2004
落叶乔木Deciduous trees	海南蒲桃 Syzygium hainanense, 糖胶树 Alstonia scholaris	当年 Current year	OTC	75	40%-50%土壤相对含水量 40%-50% Relative soil water content	光合生理 Photosynthetic physiology	Hao et al., 2014
	长芒杜英 Elaeocarpus apiculatus, 壳菜果Mytilaria laosensis,黧蒴锥 Castanopsis fissa 醉香含笑 Michelia macclurei, 樟 Cinnamomum camphora, 红花荷Rhodoleia championii, 壳菜果 Mytilaria laosensis	当年 Current year 1	OTC OTC	150 20	50%对照灌溉次数 Half the watered times to control 两天一次灌溉 Watered once in two days	荧光生理 Chlorophyll a fluorescence 根、茎、叶和总生物量及根茎比 Root biomass, stem biomass, leaf biomass, total biomass and root/shoot ratio	Li et al., 2015 Ye et al., 2014
	Fagus crenata	3	GC	60	70%对照灌溉水量 70% Water supplied to control	叶片物候和芽抗寒性 Leaf phenological characteristics and bud frost hardiness	Yonekura et al., 2004
	Fagus crenata	3	GC	60	70%对照灌溉水量 70% Water supplied to control	叶片抗氧化系统 Leaf antioxidative systems	Watanabe et al., 2005
	Fagus sylvatica	60	NE	2 × AA	NE (3年不同气候) NE (Climatic variation in three years)	生理生化 Physiology and biochemistry	Löw et al., 2006
	Fagus sylvatica	60	NE	2 × AA	NE (2年不同气候) NE (Climatic variation in two years)	细根抗氧化物 Antioxidants in fine roots	Haberer et al., 2008
	Fagus sylvatica	-	NE	AA	NE (9个不同地点) NE (Nine sites)	日生长 Daily growth	Kuehn et al., 2015
	Fagus sylvatica, 夏栎 Quercus robur, 辽杨×中东杨 Populus maximowiczii × P. berolinensis	1	OTC	0.95 × AA	50%田间持水量 50% Field capacity	生长和生理响应 Growth and physiological responses	Pollastrini et al., 2010
	岳桦 Betula ermanii	2	GC	50	土壤水势 6.19-30.96 kPa, 30.96-97.98 kPa Soil water potential 6.19-30.96 kPa, 30.96-97.98 kPa	生长、生理生化 Growth、physiology and biochemistry	Shimizu & Feng, 2007
	元宝槭 Acer truncatum	1	OTC	102-147	40%-50%田间持水量 40%-50% Field capacity	气孔响应 Stomatal response	Wen et al., 2014
	元宝槭 Acer truncatum	1	OTC	102-147	40%-50%田间持水量 40%-50% Field capacity	生长和生理 Growth and physiology	Li et al., 2015
植物类型 Plant type	植物种类 Plants species	树龄(年) Tree-age (year)	实验方法Experimental method	O₃处理 O₃ treatment (nmol·mol^-1)	干旱处理 Drought treatment	研究内容 Research content	参考文献 Reference
落叶乔木Deciduous trees	辽杨×中东杨 Populus maximowiczii × P. berolinensis	当年 Current year	OTC	0.95 × AA	150 mL水每天灌溉 150 mL of water a day	不同冠层叶绿素荧光 Chlorophyll a fluorescence along a crown	Desotgiu et al., 2012
	辽杨×中东杨 Populus maximowiczii × P. berolinensis	当年 Current year	OTC	0.95 × AA	150 mL水每天灌溉 150 mL water a day	光合、生长和同位素 Photosynthesis、growth and stable isotope	Pollastrini et al., 2013
	欧洲山杨×银白杨 Populus tremula × P. alba	当年 Current year	GC	CF + 120	SWC 35%	生理、蛋白质组 Physiological and proteomic	Bohler et al., 2013
	辽杨×中东杨 Populus maximowiczii × P. berolinensis	当年 Current year	OTC	44-53	60%, 20%对照灌溉水量 60%, 20% Water respect to control	生理生化、生长和同位素 Physiological and biochemistry、biomass and isotope	Pollastrini et al., 2014