干旱胁迫对2种梨砧木生长及叶绿素荧光特性的影响
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摘要
以杜梨(Pyrus betulaefolia Bunge.)和豆梨(Pyrus calleryana Decne.)2 a生实生苗为材料进行盆栽控水试验,对持续性梯度干旱胁迫处理下梨砧木幼苗生长、叶片光合特性及色素含量、叶绿素荧光特性的变化进行研究,为梨树苗期栽培管理和耐旱砧木的选育提供理论参考。结果表明:干旱胁迫使梨砧木新梢相对生长量下降,生长速率减缓,杜梨受胁迫的影响程度明显小于豆梨。2种砧木幼苗净光合速率(P_n)均在干旱胁迫下逐渐下降,不同程度干旱处理间下降趋势存在差异,豆梨在中度和重度干旱处理45 d时P_n值分别下降44.81%和78.04%,而杜梨则分别下降了36.8%和74.5%。叶绿素含量在干旱胁迫下呈现相似变化规律。杜梨和豆梨在干旱条件下的最大光化学效率(F_v/F_m)、实际光化学量子效率(Φ_(PSⅡ))、光化学猝灭系数(q_P)和电子传递效率(ETR)均呈下降趋势,最小初始荧光(F_0)和非光化学猝灭系数(NPQ)均显著增加,在45 d时轻度、中度、重度干旱分别使杜梨的NPQ值提高21.7%、42.7%和68.9%,而豆梨提高12.0%、42.4%和53.0%。可见,水分亏缺使梨砧木叶片叶绿素含量降低,叶片发生了光抑制,光系统Ⅱ(PSⅡ)受到破坏,光合作用明显减弱,从而抑制新梢枝条生长。但杜梨相较于豆梨能够更有效地保护光合机构,以应对胁迫对其的破坏。
The two-year-old seedlings of Pyrus betulaefolia Bunge and Pyrus calleryana Decne were used as materials to conduct potted water control tests,the change characteristics of the growth,photosynthesis,the contents of chlorophyll and carotenoid,chlorophyll fluorescence parameters were studied,this wouldlay foundation for pear seedlings cultivation and drought-tolerant rootstocks breeding.The results showed that the relative growth and the new shootsgrowth rate of pear rootstocksdecreased under drought stress,P.betulaefolia Bunge was less affected by stress than P.calleryana Decne.The net photosynthetic rate(P_n) of the two kinds of rootstocks gradually decreased under drought stress,and there were significant differences among different drought treatments.The P_n value of P.calleryana Decne decreased by 44.81% and 78.04% after 45 days under treatments of moderate and severe drought respectively,while that of P.betulaefolia Bunge decreased by 36.8% and 74.5% respectively.There was a consistency between the change trend of chlorophyll content and net photosynthetic rate.The maximum photochemical efficiency(F_v/F_m),actual photochemical quantum efficiency(Φ_(PSⅡ)),photochemical quenching coefficient(q_P) and electron transport efficiency(ETR) of both P.betulaefolia Bunge and P.calleryana Decne decreased,the minimum initial fluorescence(F_0) and non-photochemical quenching coefficient(NPQ) significantly increased.At 45 days,the NPQ value of P.betulaefolia Bunge increased by 21.7%,42.7%,68.9%,while P.calleryana Decne increased by 12.0%,42.4%,53.0% under the mild,moderate and severe drought stress respectively.Therefore,the chlorophyll content reduced,the leaves of the pear rootstocks were photoinhibitedand the photosystemⅡ(PSⅡ) was destroyed in case of water deficit,the photosynthesis of both the leaves of Pyrus betulaefolia Bunge.and Pyrus calleryana Decne seedlings were obviously weakened,and the shoot growth were inhibited.However,damages caused by stress,photosynthetic apparatus in P.betulaefolia Bunge could be protected more effectively than that in Pyrus calleryana Decne.
The two-year-old seedlings of Pyrus betulaefolia Bunge and Pyrus calleryana Decne were used as materials to conduct potted water control tests,the change characteristics of the growth,photosynthesis,the contents of chlorophyll and carotenoid,chlorophyll fluorescence parameters were studied,this wouldlay foundation for pear seedlings cultivation and drought-tolerant rootstocks breeding.The results showed that the relative growth and the new shootsgrowth rate of pear rootstocksdecreased under drought stress,P.betulaefolia Bunge was less affected by stress than P.calleryana Decne.The net photosynthetic rate(P_n) of the two kinds of rootstocks gradually decreased under drought stress,and there were significant differences among different drought treatments.The P_n value of P.calleryana Decne decreased by 44.81% and 78.04% after 45 days under treatments of moderate and severe drought respectively,while that of P.betulaefolia Bunge decreased by 36.8% and 74.5% respectively.There was a consistency between the change trend of chlorophyll content and net photosynthetic rate.The maximum photochemical efficiency(F_v/F_m),actual photochemical quantum efficiency(Φ_(PSⅡ)),photochemical quenching coefficient(q_P) and electron transport efficiency(ETR) of both P.betulaefolia Bunge and P.calleryana Decne decreased,the minimum initial fluorescence(F_0) and non-photochemical quenching coefficient(NPQ) significantly increased.At 45 days,the NPQ value of P.betulaefolia Bunge increased by 21.7%,42.7%,68.9%,while P.calleryana Decne increased by 12.0%,42.4%,53.0% under the mild,moderate and severe drought stress respectively.Therefore,the chlorophyll content reduced,the leaves of the pear rootstocks were photoinhibitedand the photosystemⅡ(PSⅡ) was destroyed in case of water deficit,the photosynthesis of both the leaves of Pyrus betulaefolia Bunge.and Pyrus calleryana Decne seedlings were obviously weakened,and the shoot growth were inhibited.However,damages caused by stress,photosynthetic apparatus in P.betulaefolia Bunge could be protected more effectively than that in Pyrus calleryana Decne.
引文
[1]安玉艳,梁宗锁,郝文芳.杠柳幼苗对不同强度干旱胁迫的生长与生理响应[J].生态学报,2011,31(3):716-725.AN Y Y,LIANG Z S,HAO W F.Growth and physiological responses of the Periploca sepium Bunge seedlings to drought stress[J].Acta Ecologica Sinica,2011,31(3):716-725.
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[7]薛杨,宋健坤,李鼎立,等.梨砧木种质资源的SSR遗传多样性分析[J].植物遗传资源学报,2013,14(6):1190-1195.XUE Y,SONG J K,LI D L,et al.SSR analysis of genetic diversity in pear rootstock germplasm[J].Journal of Plant Genetic Resources,2013,14(6):1190-1195.
[8]李迎春,樊卫国,陈双林.干旱胁迫对梨属4个重要种幼苗膜脂过氧化和抗氧化酶活性的影响[J].浙江农林大学学报,2008,25(4):437-441.LI Y CH,FAN W G,CHEN SH L.Soil drought stress on membrane-lipid peroxidation and antioxidant enzymes in pear rootstock[J].Journal of Zhejiang A&F University,2008,25(4):437-441.
[9]BANKS J M.Chlorophyll fluorescence as a tool toidentify drought stress in acer genotypes[J].Environmental and Experimental Botany,2018,155:118-127.
[10]NAIDOO G,NAIDOO K K.Drought stress effects on gas exchange and water relations of the invasive weed Chromolaena odorata[J].Flora,2018,248:1-9.
[11]麻云霞,李钢铁,张宏武,等.酸枣幼苗光合特征和生理生化指标对不同强度干旱胁迫的响应[J].干旱区资源与环境,2018,32(12):164-169.MA Y X,LI G T,ZHANG H W,et al.Photosynthetic characteristics and physiological and biochemical indexes in response to drought stress in Zizyphus jujuba seedlings[J].Journal of Arid Land Resources and Environment,2018,32(12):164-169.
[12]WANG ZH B,LI G F,SUN H Q,et al.Effects of drought stress on photosynthesis and photosynthetic electron transport chain in young apple tree leaves[J].Biology Open,2018,7(11):bio035279.
[13]曹慧,王孝威,邹岩梅,等.外源NO对干旱胁迫下平邑甜茶幼苗叶绿素荧光参数和光合速率的影响[J].园艺学报,2011,38(4):613-620.CAO H,WANG X W,ZOU Y M,et al.Membrane lipid peroxidation damage effect of chlorophyll degradation in amalus seedlings under water stress[J].Scientia Agricultura Sinica,2011,38(4):613-620.
[14]STELLFELDT A,MALDONADO M A,HUESO J J,et al.Gas exchange and water relations of young potted loquat cv.Algerie under progressive drought conditions[J].Journal of Integrative Agriculture,2018,17(6):1360-1368.
[15]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006.GAO J F.Plant Physiology Experiment Guide[M].Beijing:Higher Education Press,2006.
[16]李泽,谭晓风,卢锟,等.干旱胁迫对两种油桐幼苗生长、气体交换及叶绿素荧光参数的影响[J].生态学报,2017,37(5):1515-1524.LI Z,TAN X F,LU K,et al.Influence of drought stress on the growth,leaf gas exchange and chlorophyll fluorescence in two varieties of tung tree seedlings[J].Acta Ecologica Sinica,2017,37(5):1515-1524.
[17]ROHIT J,WANI S H,BALWANT S,et al.Transcription factors and plants response to drought stress:current under standing and future directions[J].Frontiers in Plant Science,2016,7:1029.
[18]LIN ZH F,LI SH SH,LIN G ZH,et al.Superoxide dismutase activity and lipid peroxidation in relation to senescence of rice on leaves[J].Journal of Integrative Plant Biology,1984(6):47-57.
[19]井大炜,邢尚军,杜振宇,等.干旱胁迫对杨树幼苗生长、光合特性及活性氧代谢的影响[J].应用生态学报,2013,24(7):1809-1816.JING D W,XING SH J,DU ZH Y,et al.Effects of drought stress on the growth,photosynthetic characteristics,and active oxygen metabolism of poplar seedlings[J].Chinese Journal of Applied Ecology,2013,24(7):1809-1816.
[20]LIMBU S,SHARMA L,RAO A P.Growth and photosynthetic gas exchange characteristics in Solanum aethiopicum under water stress in organic production system[J].Journal of Pharmacognosy and Phytochemistry,2018,7(2):1180-1182.
[21]FARQUHAR G D,VON CAEMMERER S,BERRY J A.Models of photosynthesis[J].Plant Physiology,2001,125(1):42-45.
[22]JALINK H,VAN DER SCHOOR R.Role of Fluorescence Approaches to Understand Functional Traits of Photosynthesis[M]//Phenomics in Crop Plants:Trends,Options and Limitations.New Delhi:Springer,2015:181-194.
[23]DIAS M C,CORREIA S,SERODIO J,et al.Chlorophyll fluorescence and oxidative stress endpoints to discriminate olive cultivars tolerance to drought and heat episodes[J].Scientia Horticulturae,2018,231:31-35.
[24]EWERS B E,PLEBAN J R,ASTON T,et al.Chlorophyll fluorescence is a rigorous,high throughput tool to analyze the impacts of genotype,species,and stress on plant and ecosystem productivity[C]//New Orleans:AGU Fall Meeting Abstracts,2017.
[25]宋丰萍,蒙祖庆,次仁德吉,等.干旱胁迫对油菜花期光合特性的影响[J].西北农业学报,2014,23(8):99-105.SONG F P,MENG Z Q,CIRENDEJI,et al.Influence of drought stress on photosynthetic traits during rapeseed flowering[J].Acta Agriculturae Boreali-occidentalis Sinica,2014,23(8):99-105.
[26]PAPAGEORGIOU G C,GOVINDJEE.Chlorophyll a Fluorescence:a Signature of Photosynthesis[M].Berlin:Springer Science&Business Media,2007.
[27]NULTSCH W,PFAU J,HUPPERTZ K.Photoinhibition of photosynthetic oxygen production and its recovery in the subtidal red algaPolyneura hilliae[J].Botanica Acta,1990,103(1):62-67.
[28]XU CH CH,LI D Q,ZOU Q,et al.Effect of drought on chlorophyll fluorescence and xanthophyll cycle components in winter wheat leaves with different ages[J].Acta Photophysiologica Sinica,1999,25(1):29-37.
[29]ANDERSSON B,ARO E M.Photodamage and D1protein turnover in photosystemⅡ[M]//Regulation of Photosynthesis.Dordrecht:Springer,2001:377-393.
[30]GODDE D,HEFER M.Photoinhibition and light-dependent turnover of the D1reaction-centre polypeptide of photosystemⅡare enhanced by mineral-stress conditions[J].Planta,1994,193(2):290-299.
[31]李耕,高辉远,赵斌,等.灌浆期干旱胁迫对玉米叶片光系统活性的影响[J].作物学报,2009,35(10):1916-1922.LI G,GAO H Y,ZHAO B,et al.Effects of drought stress on activity of photosystems in leaves of maize at grain filling stage[J].Acta Agronomica Sinica,2009,35(10):1916-1922.
[32]孙璐,周宇飞,李丰先,等.盐胁迫对高粱幼苗光合作用和荧光特性的影响[J].中国农业科学,2012,45(16):3265-3272.SUN L,ZHOU Y F,LI F X,et al.Impacts of salt stress on characteristics of photosynthesis and chlorophyll fluorescence of sorghum seedlings[J].Scientia Agricultura Sinica,2012,45(16):3265-3272.
[33]RAZAVI F,POLLET B,STEPPE K,et al.Chlorophyll fluorescence as a tool for evaluation of drought stress in strawberry[J].Photosynthetica,2008,46(4):631-633.
[2]任小云,宋宇琴,李俊豪,等.垄沟覆地布对晋西旱塬梨园土壤墒情及导水特性的影响[J].西北农业学报,2018,27(6):846-853.REN X Y,SONG Y Q,LI J H,et al.Effects of ridge-furrow polypropylene film mulching on soil moisture and hydraulic conductivity in a pear orchard in dryland highland of western Shanxi[J].Acta Agriculturae Boreali-occidentalis Sinica,2018,27(6):846-853.
[3]孙继亮,李六林,陶书田,等.干旱胁迫和复水对梨幼树生理特性的影响[J].应用与环境生物学报,2012,18(2):218-223.SUN J L,LI L L,TAO SH T,et al.Effects of drought stress and rewatering on physiological characteristics of pear seedling[J].Chinese Journal of Applied and Environmental Biology,2012,18(2):218-223.
[4]王延秀,贾旭梅,石晓昀,等.三种苹果砧木应对干旱胁迫的超微及解剖结构响应特性[J].植物生理学报,2018,54(4):594-606.WANG Y X,JIA X M,SHI X Y,et al.The response characteristics of the ultrastructure and anatomical structure of three apple rootstocks under drought stress[J].Plant Physiology Journal,2018,54(4):594-606.
[5]董星光,曹玉芬,王昆,等.中国3个主要梨砧木资源木质部导管分子结构及分布比较[J].植物学报,2015,50(2):227-233.DONG X G,CAO Y F,WANG K,et al.Comparison of the characters and distribution of vessel elements in xylem among three main pear rootstocks in China[J].Bulletin of Botany,2015,50(2):227-233.
[6]樊卫国,李迎春.部分梨砧木的叶片组织结构与抗旱性的关系[J].果树学报,2008,25(1):17-21.FAN W G,LI Y CH.Study on the relationship between lamina anatomical structure and drought resistance of pear rootstocks[J].Journal of Fruit Science,2008,25(1):17-21.
[7]薛杨,宋健坤,李鼎立,等.梨砧木种质资源的SSR遗传多样性分析[J].植物遗传资源学报,2013,14(6):1190-1195.XUE Y,SONG J K,LI D L,et al.SSR analysis of genetic diversity in pear rootstock germplasm[J].Journal of Plant Genetic Resources,2013,14(6):1190-1195.
[8]李迎春,樊卫国,陈双林.干旱胁迫对梨属4个重要种幼苗膜脂过氧化和抗氧化酶活性的影响[J].浙江农林大学学报,2008,25(4):437-441.LI Y CH,FAN W G,CHEN SH L.Soil drought stress on membrane-lipid peroxidation and antioxidant enzymes in pear rootstock[J].Journal of Zhejiang A&F University,2008,25(4):437-441.
[9]BANKS J M.Chlorophyll fluorescence as a tool toidentify drought stress in acer genotypes[J].Environmental and Experimental Botany,2018,155:118-127.
[10]NAIDOO G,NAIDOO K K.Drought stress effects on gas exchange and water relations of the invasive weed Chromolaena odorata[J].Flora,2018,248:1-9.
[11]麻云霞,李钢铁,张宏武,等.酸枣幼苗光合特征和生理生化指标对不同强度干旱胁迫的响应[J].干旱区资源与环境,2018,32(12):164-169.MA Y X,LI G T,ZHANG H W,et al.Photosynthetic characteristics and physiological and biochemical indexes in response to drought stress in Zizyphus jujuba seedlings[J].Journal of Arid Land Resources and Environment,2018,32(12):164-169.
[12]WANG ZH B,LI G F,SUN H Q,et al.Effects of drought stress on photosynthesis and photosynthetic electron transport chain in young apple tree leaves[J].Biology Open,2018,7(11):bio035279.
[13]曹慧,王孝威,邹岩梅,等.外源NO对干旱胁迫下平邑甜茶幼苗叶绿素荧光参数和光合速率的影响[J].园艺学报,2011,38(4):613-620.CAO H,WANG X W,ZOU Y M,et al.Membrane lipid peroxidation damage effect of chlorophyll degradation in amalus seedlings under water stress[J].Scientia Agricultura Sinica,2011,38(4):613-620.
[14]STELLFELDT A,MALDONADO M A,HUESO J J,et al.Gas exchange and water relations of young potted loquat cv.Algerie under progressive drought conditions[J].Journal of Integrative Agriculture,2018,17(6):1360-1368.
[15]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006.GAO J F.Plant Physiology Experiment Guide[M].Beijing:Higher Education Press,2006.
[16]李泽,谭晓风,卢锟,等.干旱胁迫对两种油桐幼苗生长、气体交换及叶绿素荧光参数的影响[J].生态学报,2017,37(5):1515-1524.LI Z,TAN X F,LU K,et al.Influence of drought stress on the growth,leaf gas exchange and chlorophyll fluorescence in two varieties of tung tree seedlings[J].Acta Ecologica Sinica,2017,37(5):1515-1524.
[17]ROHIT J,WANI S H,BALWANT S,et al.Transcription factors and plants response to drought stress:current under standing and future directions[J].Frontiers in Plant Science,2016,7:1029.
[18]LIN ZH F,LI SH SH,LIN G ZH,et al.Superoxide dismutase activity and lipid peroxidation in relation to senescence of rice on leaves[J].Journal of Integrative Plant Biology,1984(6):47-57.
[19]井大炜,邢尚军,杜振宇,等.干旱胁迫对杨树幼苗生长、光合特性及活性氧代谢的影响[J].应用生态学报,2013,24(7):1809-1816.JING D W,XING SH J,DU ZH Y,et al.Effects of drought stress on the growth,photosynthetic characteristics,and active oxygen metabolism of poplar seedlings[J].Chinese Journal of Applied Ecology,2013,24(7):1809-1816.
[20]LIMBU S,SHARMA L,RAO A P.Growth and photosynthetic gas exchange characteristics in Solanum aethiopicum under water stress in organic production system[J].Journal of Pharmacognosy and Phytochemistry,2018,7(2):1180-1182.
[21]FARQUHAR G D,VON CAEMMERER S,BERRY J A.Models of photosynthesis[J].Plant Physiology,2001,125(1):42-45.
[22]JALINK H,VAN DER SCHOOR R.Role of Fluorescence Approaches to Understand Functional Traits of Photosynthesis[M]//Phenomics in Crop Plants:Trends,Options and Limitations.New Delhi:Springer,2015:181-194.
[23]DIAS M C,CORREIA S,SERODIO J,et al.Chlorophyll fluorescence and oxidative stress endpoints to discriminate olive cultivars tolerance to drought and heat episodes[J].Scientia Horticulturae,2018,231:31-35.
[24]EWERS B E,PLEBAN J R,ASTON T,et al.Chlorophyll fluorescence is a rigorous,high throughput tool to analyze the impacts of genotype,species,and stress on plant and ecosystem productivity[C]//New Orleans:AGU Fall Meeting Abstracts,2017.
[25]宋丰萍,蒙祖庆,次仁德吉,等.干旱胁迫对油菜花期光合特性的影响[J].西北农业学报,2014,23(8):99-105.SONG F P,MENG Z Q,CIRENDEJI,et al.Influence of drought stress on photosynthetic traits during rapeseed flowering[J].Acta Agriculturae Boreali-occidentalis Sinica,2014,23(8):99-105.
[26]PAPAGEORGIOU G C,GOVINDJEE.Chlorophyll a Fluorescence:a Signature of Photosynthesis[M].Berlin:Springer Science&Business Media,2007.
[27]NULTSCH W,PFAU J,HUPPERTZ K.Photoinhibition of photosynthetic oxygen production and its recovery in the subtidal red algaPolyneura hilliae[J].Botanica Acta,1990,103(1):62-67.
[28]XU CH CH,LI D Q,ZOU Q,et al.Effect of drought on chlorophyll fluorescence and xanthophyll cycle components in winter wheat leaves with different ages[J].Acta Photophysiologica Sinica,1999,25(1):29-37.
[29]ANDERSSON B,ARO E M.Photodamage and D1protein turnover in photosystemⅡ[M]//Regulation of Photosynthesis.Dordrecht:Springer,2001:377-393.
[30]GODDE D,HEFER M.Photoinhibition and light-dependent turnover of the D1reaction-centre polypeptide of photosystemⅡare enhanced by mineral-stress conditions[J].Planta,1994,193(2):290-299.
[31]李耕,高辉远,赵斌,等.灌浆期干旱胁迫对玉米叶片光系统活性的影响[J].作物学报,2009,35(10):1916-1922.LI G,GAO H Y,ZHAO B,et al.Effects of drought stress on activity of photosystems in leaves of maize at grain filling stage[J].Acta Agronomica Sinica,2009,35(10):1916-1922.
[32]孙璐,周宇飞,李丰先,等.盐胁迫对高粱幼苗光合作用和荧光特性的影响[J].中国农业科学,2012,45(16):3265-3272.SUN L,ZHOU Y F,LI F X,et al.Impacts of salt stress on characteristics of photosynthesis and chlorophyll fluorescence of sorghum seedlings[J].Scientia Agricultura Sinica,2012,45(16):3265-3272.
[33]RAZAVI F,POLLET B,STEPPE K,et al.Chlorophyll fluorescence as a tool for evaluation of drought stress in strawberry[J].Photosynthetica,2008,46(4):631-633.