干旱条件下栓皮栎叶片光系统活性受损的温度指标
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摘要
利用盆栽控制土壤水分和气候箱控制温度的方法,定量研究高温对栓皮栎幼苗胁迫的温度指标。设定土壤体积含水量>14.5%(无干旱胁迫,T_0)、12.5%～14.5%(轻度干旱,T_1)、9.5%～11.5%(中度干旱胁迫,T_2)、5.5%～7.5%(重度干旱胁迫,T_3)4种土壤水分处理水平,在气温35、37、39、…、49℃下,各温度处理时间均设置为1、2、…、6h。通过测定干旱条件下栓皮栎叶片光系统PSⅡ最大量子效率Fv/Fm、光合量子产额Y(Ⅱ)、调节性热耗散Y(NPQ)和非调节性热耗散Y(NO)等叶绿素荧光参数随温度的变化过程,确定叶片光系统活性受损的温度指标。结果表明:(1)从T_0、T_1、T_2到T_3,Fv/Fm、Y(Ⅱ)随温度上升其曲线的降低幅度逐渐增加,说明干旱水平对胁迫温度指标有较大影响;(2)由Fv/Fm确定的不同干旱水平PSⅡ受损的温度指标为51.6～57.5℃;而由Y(Ⅱ)确定的胁迫影响光合作用产率的温度指标为40.7～52.5℃;(3)Y(NPQ)开始上升的温度由T_0、T_1的45℃降至T_2的43℃、T_3的41℃;(4)Y(NO)的变化表现为T_0不升高,T_1大于47℃时开始升高,T_2和T_3在45℃以上开始升高。说明干旱和高温叠加条件下,PSⅡ受损的温度和光合作用产率下降的温度指标与无干旱相比有较大差异,干旱胁迫越严重叶片光系统PSⅡ受损的温度指标越低。干旱使栓皮栎主动和被动热耗散均增加,主动耗散过剩光能开始增加的温度低于被动热耗散开始增加的温度。
The temperature index of Quercus variabilis seedling in high temperature and drought stress was determined using method of soil water controlling in pot and temperature controlling in climate box. Soil volumetric humidity >14.5% was set as no drought stress(T_0), 12.5%-14.5% was set as mild drought(T_1), 9.5%-11.5% was set as moderate drought(T_2) and 5.5%-7.5% was set as severe drought(T_3). The temperature was controlled at 35,37,39,…,49℃ and at any temperature of which keeping treating time as 1,2,…,6 h. The parameters of Chlorophyll Fluorescence measured were Fv/Fm, Y(Ⅱ), ETR, Y(NPQ) and Y(NO), which were used to determine stress temperature index. The results showed:(1) from mild, moderate to severe drought, the changing curves of Fv/Fm,Y(Ⅱ) adapt to temperature decreased more rapidly, which indicated the drought serious level had obvious significance on temperature stress index.(2)The temperature indices of different drought levels indicating PSⅡ damaging determined by Fv/Fm were 51.6-57.5℃; The temperature indices of different drought levels indicating PSⅡ efficiency of photosynthesis determined by Y(Ⅱ) were 40.7-52.5℃;(3) Regulatory heat dissipation Y(NPQ)began to rise at 45℃ by T_0 and T_1, and at 43℃ by T_2, 41℃ by T_3;(4) Nonregulatory heat dissipation Y(NO) of T_0 hardly rise, and T_1 began to rise at 47℃, T_2 and T_3 began to rise at 45℃. The results above indicated the temperature indices of PSⅡ damaged and photosynthetic efficiency decreased under superimposed drought conditions were significantly different from those without drought. The more drought, the lower the stress temperature index. Both active and passive heat dissipations of Quercus variabilis were increased by drought stress, and the temperature at which excess light energy of active dissipation began to increase was lower than that at which passive heat dissipation began to increase.
The temperature index of Quercus variabilis seedling in high temperature and drought stress was determined using method of soil water controlling in pot and temperature controlling in climate box. Soil volumetric humidity >14.5% was set as no drought stress(T_0), 12.5%-14.5% was set as mild drought(T_1), 9.5%-11.5% was set as moderate drought(T_2) and 5.5%-7.5% was set as severe drought(T_3). The temperature was controlled at 35,37,39,…,49℃ and at any temperature of which keeping treating time as 1,2,…,6 h. The parameters of Chlorophyll Fluorescence measured were Fv/Fm, Y(Ⅱ), ETR, Y(NPQ) and Y(NO), which were used to determine stress temperature index. The results showed:(1) from mild, moderate to severe drought, the changing curves of Fv/Fm,Y(Ⅱ) adapt to temperature decreased more rapidly, which indicated the drought serious level had obvious significance on temperature stress index.(2)The temperature indices of different drought levels indicating PSⅡ damaging determined by Fv/Fm were 51.6-57.5℃; The temperature indices of different drought levels indicating PSⅡ efficiency of photosynthesis determined by Y(Ⅱ) were 40.7-52.5℃;(3) Regulatory heat dissipation Y(NPQ)began to rise at 45℃ by T_0 and T_1, and at 43℃ by T_2, 41℃ by T_3;(4) Nonregulatory heat dissipation Y(NO) of T_0 hardly rise, and T_1 began to rise at 47℃, T_2 and T_3 began to rise at 45℃. The results above indicated the temperature indices of PSⅡ damaged and photosynthetic efficiency decreased under superimposed drought conditions were significantly different from those without drought. The more drought, the lower the stress temperature index. Both active and passive heat dissipations of Quercus variabilis were increased by drought stress, and the temperature at which excess light energy of active dissipation began to increase was lower than that at which passive heat dissipation began to increase.
引文
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[13]余凌翔,朱勇,钟楚,等.生态环境对烟草农艺性状和叶绿素荧光特性的影响[J].中国农业气象,2015,36(2):149-154.Yu L X,Zhu Y,Zhong C,et al.Effects of ecological environment on tobacco agronomic traits and chlorophyll fluorescence characteristic[J].Chinese Journal of Agrometeorology,2015,36(2):149-154.(in Chinese)
[14]杨再强,李伶俐,殷剑敏,等.灌浆初期不同时长高温胁迫对早稻叶片光合和荧光参数的影响[J].中国农业气象,2014,35(1):80-84.Yang Z Q,Li L L,Yin J M,et al.Effects of different duration of high temperature stress during filling stages on leaf photosynthetic and fluorescence parameters of early rice[J].Chinese Journal of Agrometeorology,2014,35(1):80-84.(in Chinese)
[15]陈景玲,宋晓明,王谦,等.基于叶绿素荧光参数的栓皮栎叶片PSⅡ失活高温指标[J].中国农业气象,2013,34(5):563-568.Chen J L,Song X M,Wang Q,et al.High temperature index of PSⅡinactivation according to chlorophyll fluorescence of quercus variabilis leaves[J].Chinese Journal of Agrometeorology,2013,34(5):563-568.(in Chinese)
[16]张文辉,段宝利,周建云,等.不同种源栓皮栎幼苗水分适应及耐旱特性比较研究[J].西北植物学报,2003,23(5):728-734.Zhang W H,Duan B L,Zhou J Y,et al.A comparative study on characters of drought resistance for four provenances of Quercus variabilis[J].Acta Botanica Boreali-Occidentalia Sinica,2003,23(5):728-734.(in Chinese)
[17]Ulrich Schreiber.叶绿素荧光基础实验指南[M].韩志国译.上海:德国WALZ公司中国技术服务中心,2004:23-24.Ulrich Schreiber.Chlorophyll fluorescence and photosynthetic energy conversion:simple introductory experiments with the TEACHING-PAM Chlorophyll Fluorometer[M].Translated by Han Z G.Heinz Walz GmbH,2004:23-24.(in Chinese)
[18]王莹博,许申平,马杰,崔波.高温干旱胁迫对白及光合特性及叶绿素荧光参数的影响[J].河南农业大学学报,2018,52(2):199-205.Wang Y B,Xu S P,Ma J,et al.Effects of heat drought stress on the photosynthetic characteristics and chlorophyll fluorescence parameters of Bletillastriata[J].Journal of Henan Agricultural University,2018,52(2):199-205.(in Chinese)
[19]赵薇,李贞,王谦,等.冬小麦灌浆期旗叶光曲线及叶绿素荧光参数研究[J].气象与环境科学,2017,40(1):64-72.Zhao W,Li Z,Wang Q,et al.Study on flag leaf light curve and chlorophyll fluorescence parameter of winter wheat in grain filling stage[J].Meteorological and Environmental Sciences,2017,40(1):64-72.
[20]魏丹丹,张劲松,孟平,等.栓皮栎人工林冠层温度变化特征及其与微气象因子的关系[J].应用生态学报,2012,23(7):1767-1773.Wei D D,Zhang J S,Meng P,et al.Variations of canopy temperature in Quercus variabilis plantation and their relations with micrometeorological factors[J].Chinese Journal of Applied Ecology,2012,23(7):1767-1773.(in Chinese)
[21]韩旖旎.两种沙生灌木适应波动环境的非光化学淬灭调节[D].北京:北京林业大学,2016.Han Y N.Regulation of non-photochemical quenching in photosynthetica acclimation of two xerphytic shrubs to fluctuating environments[D].Beijing:Beijing Forestry University,2016.(in Chinese)
[22]张金恩,肖洪,郑有飞,等.开顶式气室内外冬小麦光合特性差异比较[J].生态学报,2015,35(21):6993-7002.Zhang J E,Xiao H,Zheng Y F,et al.Comparative of the photosynthetic characteristics of winter wheat grown inside and outside open-top chambers[J].Acta Ecologica Sinica,2015,35(21):6993-7002.(in Chinese)
[23]何静丹.低温胁迫及温度回升对甘蔗生理生化指标的影响[D].南宁:广西大学,2011.He J D.Effects of low temperature stress and temperature rewarming on the physiological and biochemical indexes of sugarcane[D].Nanning:Guangxi Unversity,2011.(in Chinese)
[24]张金恩.UV-B辐射增强与O3胁迫单因子及其复合作用下冬小麦光合能力变化研究[D].南京:南京信息工程大学,2011.Zhang J E.Photosynthetic capacity changes of winter wheat under UV-B radiation intensification and O3 stress and their combined effects[D].Nanjing:Nanjing University of Information Science&Technology,2011.(in Chinese)
[2]Matusick G,Ruthrof K X,Hardy G S J.Drought and heat triggers sudden and severe dieback in a dominant mediterranean-type woodland species[J].Open Journal of Forestry,2012,2(4):183-186.
[3]Behrangi A,Loikith P C,Fetzer E J,et al.Utilizing humidity and temperature data to advance monitoring and prediction of meteorological drought[J].Climate,2015,3:999-1017.
[4]Sun B F,Zhao H,Wang X K.Effects of drought on net primary productivity:roles of temperature,drought intensity,and duration[J].Chin.Geogra.Sci.,2016,26(2):270-282.
[5]Allen C D,Breshears D D,McDowell N G.On underestimation of global vulnerability to tree mortality and forest die-off from hotter drought in the Anthropocene[J].Ecosphere,2015,6(8):129.
[6]Zavalloni C,Gielen B,Boeck H J D,et al.Greater impact of extreme drought on photosynthesis of grasslands exposed to a warmer climate in spite of acclimation[J].Physiologia Plantarum,2009,136:57-72.
[7]Wardlaw I F.Interaction between drought and chronic high temperature during kernel filling in wheat in a controlled environment[J].Annals of Botany,2002,90:469-476.
[8]Prasad P V V,Pisipati S R,Mom?ilovi?I,et al.Independent and combined effects of high temperature and drought stress during grain filling on plant yield and chloroplast EF-Tu expression in spring wheat[J].Journal of Agronomy and Crop Science,2011,197(6):430-441.
[9]Wang G P,Tian F X,Zhang M,et al.The over accumulation of glycinebetaine alleviated damages to PSII of wheat flag leaves under drought and high temperature stress combination[J].Acta Physiol Plant,2014,36:2743-2753.
[10]秦立琴,张悦丽,郭峰,等.强光下高温与干旱胁迫对花生光系统的伤害机制[J].生态学报,2011,31(7):1835-1843.Qin L Q,Zhang Y L,Guo F,et al.Damaging mechanisms of peanut(Arachis hypogaea L.)photosystems caused by high-temperature and drought under high irradiance[J].Acta Ecologica Sinica,2011,31(7):1835-1843.(in Chinese)
[11]李凯伟,杨再强,肖芳,等.寡照胁迫对设施葡萄叶片光合特性的影响及评价[J].中国农业气象,2017,38(12):801-811.Li K W,Yang Z Q,Xiao F,et al.Effects and evaluation of low irradiation stress on photosynthetic characteristics of grapevine leaves in greenhouse[J].Chinese Journal of Agrometeorology,2017,38(12):801-811.(in Chinese)
[12]熊宇,杨再强,薛晓萍,等.遮光处理对温室黄瓜幼龄植株叶片光合参数的影响[J].中国农业气象,2016,37(2):222-230.Xiong Y,Yang Z Q,Xue X P,et al.Effect of shading on photosynthetic parameters in greenhouse cucumber leaves[J].Chinese Journal of Agrometeorology,2016,37(2):222-230.(in Chinese)
[13]余凌翔,朱勇,钟楚,等.生态环境对烟草农艺性状和叶绿素荧光特性的影响[J].中国农业气象,2015,36(2):149-154.Yu L X,Zhu Y,Zhong C,et al.Effects of ecological environment on tobacco agronomic traits and chlorophyll fluorescence characteristic[J].Chinese Journal of Agrometeorology,2015,36(2):149-154.(in Chinese)
[14]杨再强,李伶俐,殷剑敏,等.灌浆初期不同时长高温胁迫对早稻叶片光合和荧光参数的影响[J].中国农业气象,2014,35(1):80-84.Yang Z Q,Li L L,Yin J M,et al.Effects of different duration of high temperature stress during filling stages on leaf photosynthetic and fluorescence parameters of early rice[J].Chinese Journal of Agrometeorology,2014,35(1):80-84.(in Chinese)
[15]陈景玲,宋晓明,王谦,等.基于叶绿素荧光参数的栓皮栎叶片PSⅡ失活高温指标[J].中国农业气象,2013,34(5):563-568.Chen J L,Song X M,Wang Q,et al.High temperature index of PSⅡinactivation according to chlorophyll fluorescence of quercus variabilis leaves[J].Chinese Journal of Agrometeorology,2013,34(5):563-568.(in Chinese)
[16]张文辉,段宝利,周建云,等.不同种源栓皮栎幼苗水分适应及耐旱特性比较研究[J].西北植物学报,2003,23(5):728-734.Zhang W H,Duan B L,Zhou J Y,et al.A comparative study on characters of drought resistance for four provenances of Quercus variabilis[J].Acta Botanica Boreali-Occidentalia Sinica,2003,23(5):728-734.(in Chinese)
[17]Ulrich Schreiber.叶绿素荧光基础实验指南[M].韩志国译.上海:德国WALZ公司中国技术服务中心,2004:23-24.Ulrich Schreiber.Chlorophyll fluorescence and photosynthetic energy conversion:simple introductory experiments with the TEACHING-PAM Chlorophyll Fluorometer[M].Translated by Han Z G.Heinz Walz GmbH,2004:23-24.(in Chinese)
[18]王莹博,许申平,马杰,崔波.高温干旱胁迫对白及光合特性及叶绿素荧光参数的影响[J].河南农业大学学报,2018,52(2):199-205.Wang Y B,Xu S P,Ma J,et al.Effects of heat drought stress on the photosynthetic characteristics and chlorophyll fluorescence parameters of Bletillastriata[J].Journal of Henan Agricultural University,2018,52(2):199-205.(in Chinese)
[19]赵薇,李贞,王谦,等.冬小麦灌浆期旗叶光曲线及叶绿素荧光参数研究[J].气象与环境科学,2017,40(1):64-72.Zhao W,Li Z,Wang Q,et al.Study on flag leaf light curve and chlorophyll fluorescence parameter of winter wheat in grain filling stage[J].Meteorological and Environmental Sciences,2017,40(1):64-72.
[20]魏丹丹,张劲松,孟平,等.栓皮栎人工林冠层温度变化特征及其与微气象因子的关系[J].应用生态学报,2012,23(7):1767-1773.Wei D D,Zhang J S,Meng P,et al.Variations of canopy temperature in Quercus variabilis plantation and their relations with micrometeorological factors[J].Chinese Journal of Applied Ecology,2012,23(7):1767-1773.(in Chinese)
[21]韩旖旎.两种沙生灌木适应波动环境的非光化学淬灭调节[D].北京:北京林业大学,2016.Han Y N.Regulation of non-photochemical quenching in photosynthetica acclimation of two xerphytic shrubs to fluctuating environments[D].Beijing:Beijing Forestry University,2016.(in Chinese)
[22]张金恩,肖洪,郑有飞,等.开顶式气室内外冬小麦光合特性差异比较[J].生态学报,2015,35(21):6993-7002.Zhang J E,Xiao H,Zheng Y F,et al.Comparative of the photosynthetic characteristics of winter wheat grown inside and outside open-top chambers[J].Acta Ecologica Sinica,2015,35(21):6993-7002.(in Chinese)
[23]何静丹.低温胁迫及温度回升对甘蔗生理生化指标的影响[D].南宁:广西大学,2011.He J D.Effects of low temperature stress and temperature rewarming on the physiological and biochemical indexes of sugarcane[D].Nanning:Guangxi Unversity,2011.(in Chinese)
[24]张金恩.UV-B辐射增强与O3胁迫单因子及其复合作用下冬小麦光合能力变化研究[D].南京:南京信息工程大学,2011.Zhang J E.Photosynthetic capacity changes of winter wheat under UV-B radiation intensification and O3 stress and their combined effects[D].Nanjing:Nanjing University of Information Science&Technology,2011.(in Chinese)