郝玉杰,武林楠,冯建荣,白玛多吉,王富霞,荣新民
(1. 石河子大学农学院/特色果蔬栽培生理与种质资源利用兵团重点实验室,新疆石河子 832003;2. 石河子农业科学研究院葡萄研究所/葡萄北疆试验站,新疆石河子 832000)
NaCl胁迫对两个葡萄品种叶片光合特性的影响
郝玉杰1,武林楠1,冯建荣1,白玛多吉1,王富霞2,荣新民2
(1. 石河子大学农学院/特色果蔬栽培生理与种质资源利用兵团重点实验室,新疆石河子 832003;2. 石河子农业科学研究院葡萄研究所/葡萄北疆试验站,新疆石河子 832000)
【目的】研究NaCl胁迫对两个葡萄新品种火洲红玉、火洲黑玉光合特性的影响,为筛选耐盐新品种提供理论依据。【方法】采用水培法,对两个葡萄品种进行不同浓度NaCl胁迫(0、2、3、4 g/L)处理,8 d后测定叶片中的叶绿素相对含量(SPAD)值、初始荧光(Fo)、最大荧光(Fm)、PSⅡ实际光化学效率(ФPSⅡ)、PSⅡ最大光化学效率(Fv/Fm)、净光合速率(Pn)、气孔导度(Gs)、胞间二氧化碳浓度(Ci)和蒸腾速率(Tr)。【结果】在NaCl≤3 g/L时,两品种SPAD值较对照显著升高,至4 g/L时,SPAD值均下降,火洲红玉SPAD值显著低于对照;随NaCl浓度升高,火洲黑玉的Fo值升高但较对照无显著性差异,Fm、ФPSⅡ、Fv/Fm值呈下降趋势但均与对照无显著性差异;火洲红玉Fo值在NaCl≥3 g/L时较对照显著升高,Fm、ФPSⅡ、Fv/Fm值较对照显著下降;火洲黑玉Ci在2 g/L时,较对照显著降低,在NaCl≥3 g/L时Ci升高但与对照无显著性差异,Gs在各处理下较对照均无显著性差异,Pn和Tr较对照均显著降低;火洲红玉Ci均显著高于对照,Gs、Pn、Tr均显著低于对照。【结论】火洲黑玉在4 g/L NaCl处理时,SPAD值较对照显著下降,Pn、Tr较对照显著降低,其他指标均与对照无显著性差异,火洲红玉在NaCl≥3 g/L时各项指标均与对照存在显著差异,火洲黑玉耐盐性较火洲红玉好。
葡萄;盐胁迫;光合特性;叶绿素荧光
【研究意义】土壤盐渍化已经成为一个世界性的生态问题,园艺作物生产也深受其害[1]。盐害往往对作物造成渗透胁迫、离子毒害、营养不平衡等危害,从而影响植物光合作用,抑制植物的生长,甚至导致其死亡[2]。新疆盐碱地占总耕地的31%[3],葡萄作为在新疆种植面积较大的树种之一[4],盐害已成为影响葡萄生产的主要问题之一[5-7]。因此,研究葡萄的耐盐性,对培育、推广适宜盐碱地种植的葡萄品种有重要的意义。【前人研究进展】前人对葡萄在盐胁迫下光合特性做了许多研究,如秦岭等[8]认为在100 mmol/L的NaCl胁迫下叶绿素含量降低,最终导致PSⅡ潜在活性中心受损,进而影响CO2的固定和同化,光能转化效率和净光合效率明显降低。李学孚等[9]研究表明,在NaCl>0.4%时,‘鄞红’葡萄叶绿素含量、叶绿素荧光特性、气体交换参数均呈现下降趋势。但马帅等[10]研究显示,随盐浓度的升高,葡萄的气体交换参数除胞间CO2浓度(Ci)逐渐上升外,其余参数均呈现下降的趋势。邢庆振等[11]对‘红地球’研究表明,随盐胁迫的加剧,光合特性中除Ci呈现先下降后上升趋势,其余各项指标均呈现下降趋势。秦红艳等[12]认为,随盐浓度的升高山葡萄初始荧光(Fo)显著升高,其他荧光参数均显著降低,PSⅡ反应中心被破坏。【本研究切入点】研究选取两个葡萄新品种火洲黑玉和火洲红玉,采用水培方法,从幼苗光合作用和叶绿素荧光动力学特征入手,探究其对不同浓度NaCl胁迫对两品种的叶绿素相对含量、叶绿素荧光参数及光合特性影响。【拟解决的关键问题】测定两个品种在盐胁迫下光合指标,了解两个葡萄品种光合特性对盐胁迫的响应,为两个葡萄品种在北疆地区进一步培育和推广提供理论依据。
1.1 材 料
参试的2个葡萄品种火洲黑玉,火洲红玉是吐鲁番葡萄研究所选育的欧亚种,由石河子葡萄研究所引入的2年生苗。2015年5月30日,选取生长健壮的火洲黑玉和火洲红玉葡萄幼苗各36株,从田间移出,用清水冲洗根,先置于塑料桶(上口径20 cm,下口径15 cm,高25 cm)用清水中培养1 d,用1/2Hoagland营养液[13]培养4 d,最后用全营养液进行培养,每个塑料桶中3株,作为1个重复,并用苯板固定,3次重复,培养51 d,于7月21日开始进行盐胁迫试验。
1.2 方 法
1.2.1 试验材料
设3个NaCl浓度梯度,对照为Hoagland营养液;处理Ⅰ为Hoagland营养液+2 g/L NaCl;处理Ⅱ为Hoagland营养液+3 g/L NaCl;处理Ⅲ为Hoagland营养液+4 g/L NaCl。试验期间营养液用充气泵连续通气,每天补充塑料桶中散失的水分至5 L刻度线,并且每4 d更换一次营养液。盐胁迫第8 d,从每个重复中随机选取1株,共选3株,每株从顶部向下数第3~4片功能叶用来测定叶片的叶绿素相对含量、气体交换参数及叶绿素荧光参数。
1.2.2 测定项目1.2.2.1 叶绿素含量
利用SPAD-502+PLUS叶绿素仪参照李学孚等[8]方法于10:00~12:00对叶绿素含量进行测定。
1.2.2.2 叶绿素荧光参数
选取健壮植株,使用FMS-2脉冲调制式荧光仪参照邢庆振等[11]方法,于10:00~12:00测定荧光参数,光适应下,打开内源光化光(600 μmol/(m2·s))测定光适应后最小荧光(Fs)、最大荧光(Fm’)、PSⅡ实际光化学效率ФPSⅡ,暗适应30 min后先照射检测光(<0.05 μmol/(m2·s)),测定初始荧光(Fo)、再照射饱和脉冲光(12 000μmol/(m2·s)),测定 PSⅡ最大光化学量子效率Fv/Fm。
1.2.2.3 光合指标
利用LI-6400便携式光合作用测定系统参照徐焕文等[14]方法于10:00~12:00测定光和参数,净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、胞间CO2浓度(Ci)由仪器自动给出[14]。测定时温度为(27±2)℃,光照强度为1 300 μmol/(m2·s),CO2浓度(380±10) μmol/mol。
1.3 数据统计
试验原始数据处理和绘图采用Excel软件完成,差异显著分析采用SPSS 19.0软件。
2.1 盐胁迫对两个葡萄品种叶片中叶绿素相对含量的影响
研究表明,不同浓度NaCl处理后,火洲黑玉和火洲红玉的叶绿素相对含量(SPAD)值随着NaCl浓度升高呈现先升高后降低的趋势。在NaCl≤3 g/L时两品种SPAD值较对照均显著升高且在3 g/L时达到峰值,分别为31.2和28.2,在4 g/L处理时,火洲黑玉SPAD值下降但与对照差异不显著,而火洲红玉SPAD值较对照显著降低。盐胁迫下火洲黑玉的SPAD值显著高于火洲红玉。图1
图1 盐胁迫下葡萄叶片中叶绿素相对含量变化
Fig.1 Effects of NaCl stress on the chlorophyll relative content in grape leaves
2.2 盐胁迫对不同葡萄品种叶片中荧光参数的影响
研究表明,随着NaCl浓度的升高,两个葡萄品种Fo呈现上升趋势,Fm、ФPSⅡ、Fv/Fm均呈现下降趋势。火洲黑玉Fo值在各处理时较对照无显著性差异;火洲红玉在NaCl≥3 g/L时Fo值较对照均显著升高,分别升高了55.6%、79.5%。火洲黑玉Fm值在各处理时较对照无显著性差异;火洲红玉在NaCl≥3 g/L时Fm值较对照显著下降,分别下降了38.2%和44%。火洲黑玉在各处理时ФPSⅡ值较对照无显著性差异;火洲红玉在NaCl≥3 g/L时ФPSⅡ值较对照显著下降,分别下降了14.4%和19.3%。火洲黑玉Fv/Fm值在各处理时较对照无显著性差异;火洲红玉在NaCl≥3 g/L时Fv/Fm值较对照显著下降,分别下降了14.5%和23.2%。火洲黑玉Fo值上升幅度显著低于火洲红玉,火洲黑玉Fm、ФPSⅡ、Fv/Fm值下降幅度显著低于火洲红玉,火洲红玉在NaCl≥3 g/L处理时较火洲黑玉PSⅡ受损程度更高。图2
图2 盐胁迫下葡萄叶片荧光参数变化
Fig. 2 Effects of NaCl stress on fluorescence parameters of grape leaves
2.3 盐胁迫对两个葡萄品种叶片气体交换参数的影响
研究表明,随NaCl浓度的升高两品种Pn、Tr、Gs均呈下降趋势,火洲黑玉Ci先下降后上升,而火洲红玉Ci则呈上升趋势。火洲黑玉和火洲红玉在各处理下Pn值较对照均显著降低,分别降低了12.8%、25.6%、43.6%和21.4%、42.9%、61.9%。火洲黑玉和火洲红玉Tr值在各个处理时较对照均显著降低,分别降低了22.7%、57.1%、64.2%和32.3%、44.6%、67.5%。火洲黑玉在各处理下Gs值与对照相比均未达到显著水平,较对照分别降低了10.6%、12.9%、14.8%;火洲红玉在各处理下Gs值均显著低于对照,分别降低了28.1%、24.3%、50.6%。火洲黑玉在2 g/L时Ci值显著低于对照,下降幅度为19.2%,在3 g/L、4 g/L时Ci较对照相比上升幅度未达显著水平;火洲红玉的Ci值在2、3 g/L时与对照相比无显著性差异,在4 g/L时Ci值较对照显著升高,升高了35.4%。火洲黑玉的Pn、Tr、Gs下降幅度显著低于火洲红玉。图3
图3 盐胁迫下葡萄叶片气体交换参数变化
Fig.3 Effects of NaCl stress on gas exchange parameters in grape leaves
植物叶片中叶绿素含量是衡量植物抗逆性的一个重要生理指标[15]。李学孚等[8]对‘鄞红’葡萄光合特性研究结果表明,低盐胁迫下(NaCl≤0.4%),SPAD值无显著变化,而高盐胁迫下(NaCl>0.4%)SPAD值显著下降,盐浓度(NaCl=0.4%)成为‘鄞红’葡萄耐盐性的临界阈值。试验研究结果表明,火洲黑玉和火洲红玉在(NaCl≤0.3%)时,SPAD值均表现升高的趋势,在(NaCl>0.3%)时,两个葡萄品种SPAD值表现下降的趋势,盐浓度(NaCl=0.3%)成为两品种耐盐性的临界阈值,这与李学孚等[8]研究结果不完全一致,李学孚[8]使用的‘鄞红’葡萄是欧美杂交种群,而试验使用的火洲黑玉和火洲红玉是由吐鲁番葡萄研究所选育的欧亚种,说明试验利用的火洲黑玉和火洲红玉的耐盐性没有欧美杂交的葡萄品种强。
NaCl胁迫导致植物光合作用的降低既有气孔因素,也有非气孔因素[18]。根据Farquhar等[19]提出的光合作用气体交换模型中,若光合速率的下降并且Ci的上升,那么影响光合作用的主要限制因素为非气孔因素,若光合速率下降伴随着Ci同时下降,那么主要的限制光合作用的因素则为气孔因素。试验研究表明,2 g/L处理下火洲黑玉葡萄叶片Pn、Gs下降,且Ci呈下降趋势(3-D),3 g/L、4 g/L处理下Pn和Gs下降的同时Ci呈现上升的趋势,表明在低盐浓度(NaCl<3 g/L)胁迫时气孔限制成为降低光合作用的主要因素,而高浓度(NaCl≥3 g/L)胁迫时则转为非气孔限制所致。房玉林等[20]对8804品系葡萄的研究认为随盐浓度升高葡萄光合作用的影响逐渐从气孔限制转变为非气孔限制。试验与房玉林等[19]研究结果一致。火洲红玉的Pn和Gs在整个胁迫过程中均呈现下降的趋势,Ci呈现上升趋势,表明火洲红玉对盐胁迫较为敏感,其光合作用下降都由非气孔限制所致。这与马帅等[10]对‘摩尔多瓦’研究结果一致。
在NaCl≤3 g/L处理下火洲黑玉和火洲红玉,SPAD值显著升高,NaCl>3 g/L处理下,SPAD值均下降,火洲红玉SPAD值较对照显著下降;NaCl胁迫下,两品种Fo值上升、Fm、Fv/Fm和ФPSⅡ值下降,但火洲黑玉在各处理下较对照均无显著性差异,火洲红玉在NaCl≥3 g/L处理下较对照均存在显著性差异;火洲黑玉NaCl胁迫下,Gs较对照无显著性差异,Pn、Tr及2 g/L时Ci较对照显著下降,在NaCl≥3 g/L时,Ci上升但与对照无显著性差异;火洲红玉Ci较对照显著上升,Pn、Gs、Tr均较对照显著下降。火洲红玉各项指标变化幅度均显著高于火洲黑玉,火洲黑玉耐盐性较火洲红玉好。
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Fund project:Special funds for construction projects of modern agricultural industry technology system(CARS-30)
Effects of NaCl Stress on Photosynthetic Characteristics of Leaves of the Two Grape Cultivars
HAO Yu-jie1,WU Lin-nan1,FENG Jian-rong1,Baimaduoji1,WANG Fu-xia2,RONG Xin-min2
(1.CollegeofAgronomy,ShiheziUniversity/XPCCKeyLaboratoryofSpecialFruitsandVegetablesCultivationPhysiologyandGermplasmResourcesUtilization,ShiheziXinjiang832003,China; 2GrapeTestStationofGrapeResearchInstituteofShiheziAcademyofAgriculturalSciences,ShiheziXinjiang832000,China)
【Objective】 The impact of salt stress on the photosynthetic characteristics of two new grape cultivars 'Huozhouheiyu' and 'Huozhouhongyu' were studied for providing a foundation for screening of new salt tolerant cultivars.【Method】Two cultivars in hydroponic cultures were treated with different concentrations of NaCl(0,2 g/L, 3 g/L, 4 g/L), and Chlorophyll content (SPAD), the minimal fluorescence (Fo), the maximal fluorescence(Fm), actual photochemical efficiency of PSⅡ(ФPSⅡ), the photochemical efficiency of PSⅡ(Fv/Fm), net photosynthetic rate (Pn), transpiration rate (Tr), stomatal conductance (Gs) and Intercellular CO2concentration (Ci) were determined after eight days.【Result】SPADof two cultivars were significantly higher in the 3 g/L and 4 g/L treatment than in the control treatment. In the 4 g/L treatment,SPADof two cultivars decreased, andSPADof 'Huozhouhongyu' was significantly lower than that of the control treatment.Foof 'Huozhouheiyu' increased and had no significant difference than control treatment.Fm, ФPSⅡ,Fv/Fmof 'Huozhouheiyu' decreased and had no significant difference than that of the control treatment.Foof 'Huozhouhongyu' was significantly higher in the 3 g/L and 4 g/L treatment than in the control treatment, andFm, ФPSⅡ,Fv/Fmof 'Huozhouhongyu' was significantly lower in the 3 g/L and 4 g/L treatment than in the control treatment.Ciof 'Huozhouheiyu' was significantly lower in the 2 g/L treatment than in the control treatment.Ciof 'Huozhouheiyu' increased and was not significantly different than that of the control treatment.Gsof 'Huozhouheiyu' was not significantly different than that of the control treatment.PnandTrwere significantly lower than that of the control treatment.Ciof 'Huozhouhongyu' was significantly higher than that of the control treatment, andPn,Tr,Gswere significantly lower.【Conclusion】In the 4 g/L treatment,SPADof 'Huozhouheiyu' decreased.PnandTrwere significantly lower than those of the control treatment. Other indexes were not significantly different than those of the control treatment. In the 3 g/L and 4 g/L, all indexes of 'Huozhouhongyu' were not siginificantly different than those of the control treatment. 'Huozhouhongyu' was affected more seriously than 'Huozhouheiyu' by salt stress.
grapes; salt stress; photosynthetic characteristics; chlorophyll fluorescence
10.6048/j.issn.1001-4330.2016.10.004
2016-05-12
现代农业产业技术体系建设专项资金项目(CARS-30)
郝玉杰(1993-),女,新疆人,硕士研究生,研究方向为果树种质资源与遗传育种,(E-mail)1275185994@qq.com
冯建荣(1969-),女,新疆人,教授,博士,研究方向为果树种质资源与分子辅助育种,(E-mail)fengjr102@126.com
S663.1
A
1001-4330(2016)10-1794-07